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

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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. salllla 4.83 2.89 1.70 5.72 053 1.25 1.64 (1.15) 0.20 0. 14 0.08 0.60 0.04 am 0.09 T. 'Ylril'gtlla 6.81 4. 27 1.69 8.93 0.56 UO 1.44 (17,226) 0.91 1.1 8 0.32 2.02 0.07 0.11 0.15

& Sada 1987). Digitizing was done using pendix), diagnosis, description, remarks, CONCH (methodology described in Chap­ type locality, di stribution and habitat, man et aI. , in prep) and a GTCO Micro­ elymology, and remarks. Asterisked (*) lots Digi Pad 12 x 121inked to a KA YPRO 2000 were used for shell measurements. Descrip­ microcom[luler. tions emphasize diMineti ve features from Descriptive statistics for all morpholog­ shell, radula, body pigmentation, and re­ ical variables were obtained for each pop­ productive systems of both scxes. ulation and species, and a selection of these are summarized (by species) in Tables I and Family Hydrobiidae Trosc hel, 1857 2. Discriminanl analys is (DA) incorporated Genus Pyrgulopsis Call & Pilsbry, 1886 all of the shell morphometric parameters Pyrgillopsis aardahli. new species and were constructed using all specimens Benton Valley springsnail measured for each species (considered as Figs. 10-14 separate groups). All computations were performed using SYST AT (Wilkinson 1986) M aterial examined. -Califomia: Mono Co.; Springat Bramlelle Raneh (87), USNM on an IBM ·AT. 860406 (holotype), *85 7951 (paratypes). 21 Apr 1987. Syslemalics Diagl1O.fis. -A tarae-sized species wit h Generic diagnoses are in Hershler & Sada broadly conical shell. Pen is wi th elongate (1987). Species accounts incl ude common filam ent, very small lobe. and ventral swell ­ name, sy nonomy, material examined (lo­ ing. Penial glandular ri dges, 1-3. calities numbered lin parentheses) as in Ap- Description. -Shell (Fig. 10) 2.6-3.4 mm ,so PROCEEDINGS OF T HE BIOLOGICAL SOCIETY O F WASHINGTON

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...- I" I N ", ,. •, ", ", Fig_ 2_ Sampling localities in SQuthem Owens Val­ ley and "''''t~m Indian Wells Valley. Solid lines indi· fig. 3. Sampling localities in nOll hem Owens Val­ calc se l ec t ~ d elevation contours (lif,hl) and modem ley. drainage (dark; stipple indicates lakes); dashed linC$ indicate historic drainage. including dry lake beds_ dusted or unpigmented. T entacles unpig­ mented except for dark ring along bases. high, height/ width, 140-170%. Whorls, Brown-black subepithelial pigmcnl gran­ 3. 75-4.25, moderately convex, moderately ules sometimes fonning dark band along shouldered. Body whorl inflated, heigh t 76- posterior edge of " ncrk." 82% of that of shell. Aperture ovate, aper­ Radular (Fig. 12) formula: 5-1-511-1 , tural pla:"!e near-parallcl to coiling axis (Fig. 2(3)-1-3, 22-24, 28-32 (from para types). 10c). Inner lip thickened, slightly reflected, Central tooth broadly Irapczoidal; basal adnate to small portion of or slighlly sep­ process moderately excavated. Penis (Fig. arated from body whorl. Outer lip straight, 13b-e) large (extending beyond mantic col­ thin. Umbilicus moderately open. lar), Ihin, considerably longer lhan wide. Dark, grey-black epithelial (melanic) pig­ Filament slender, moderate in length. Lobe ment on most of snout (to JUSt posterior of reduced or absent, with blum distal edge. cephalic tentacles), along anterior edge of Large, clongale glandular ridge borne on foot. on operculigcrous lobe (Fig. II). Cen­ elongate swelling of ventral penial surface. tral portions of sides of head/ foot lightly Single, smaller ridges sometimes found on VOLUME 102. NUMBER I '"

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Fig. 4. Sampling localities in Mono. long, and Adobe V alle~.

Fig. 5. Sampling localilies in Saline. Eureka, and tip of lobe, and on dorsal penial surface Deep Springs Valleys. proximal to base of filamenl. Filament darkly pigmented with subepithelial gran­ Pyrgulopsis amargosae, new species ules (Fig. J 3b). Capsule gland sub-equal to Amargosa springsnail albumen gland (Fig. 13a). Seminal recep­ Figs. IS, 16, 17a, b, 18, 19 tacle minute, positioned entirely anterior 10 bursa copulatrix. Bursa pear-shaped, mod­ Material Examined. -California: San erate in sizc relative to pall ial oviduct. po­ Bernardino Co.; Saratoga Spring (204), sitioned panly posterior to al bumen gla nd. USNM 860401 (hoIOlype), "8,S3,Sl's (para­ Type locality. -Spring at Bra mlette Ranch types), 7 Feb 1985; 857972, 10 l ui 1986.­ in Benton Valley, Mono County, California. Inyo Co.; Spring crossi ng path on south side Distribution and habita/.-Endemic to of Amargosa Gorge (208), *853516, 13 Mar type localilY (Fig. 14). Common in dense 1986; 857973,16 Mar 1987.-Spring feed­ watercress in uppermost portion of outflow ing Amargosa Ri ver in Amargosa Gorgc of small, hi ghly degraded spring (2O"C, 190 (208),8535 17, 13 Mar 1985.-Spring in mieromhoslcm). marsh eaSI of Grimshaw Lake (209). Etymology. - Named after Jeffrey B. Aar­ *8 53518,13 Mar 1985. dahl, Bureau of Land Management (Cali­ Diagnosis. -A small-sized species with fornia Desert District), who provided di­ globose to low conical shell. Penis large rel­ verse forms of assistance throughout the ative to head/ foot: penial lobe reduced: fil­ tenure of this project. ament long, wi de. Penial glandular ri dges, Remarks.-Distineuishcd from si milar P. 0-2. perturbata by shell appearance and hi ghly Description.-Shell (Fig. 15) 1.5-2.7 mm reduced penial lobe. high, heighllwidth, 110-160%. Whorls. 3.5- '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ,,' 17a. h. 18b-e) thickened, longe r than wide. Filament much longer than lobe, about equal 10 remaining penis le ngth. Lobe narrow, vari able in shape (Fig. 18b-e). Filament having highly muscular appearance; surface li ned wi lh weak sirialions. One or two small glandular ridges sometimes present al tip of lobe. Dark, grey-black subcpilheli al pig­ ment filling filamenllenglh al m051 10 distal tip as well as portions of remaining distal . ~ penis (Fig. JS b). Albumen and capsule glands about equal in length (Fig. 18a). Sem ­ inal receptacle minute, positioned well an· teri or 10 bursa copulalrix. Bursa copulatrix club-shaped, sma ll , positioned partly pos­ teri or to albumen gland. Type locality. -Saratoga Spri ng, southern Death Valley, San Bernard ino Co .. Califor­ nia. Distribution a"d habitat. - Found in a few .... , near-brackish springs along lower course of Amargosa River at Saratoga Spri ng, Amar­ I ' gosa Gorge, and Grimshaw Lake area (Fig. t ", " 19). Type locality an extensive marsh area (wi th several large, open pools) fed by sev­ eral small springs. Snails were common in Fi&. 6. Samphnllocalui~ In Panamlnl and nOrlh· westernmost spring, both on rocks and sub­ em IndIan Wells Valleys. mergent macrophytes in small springpool (28"C, 5000 micromhoslcm). and on emer­ 4,25, convex, moderately shouldered. Body ge nt sedges and firm bottom in shallow whorl 74-85% of shell height. Apex. pans stream outfl ow. At other localities. consist­ of teleoconch whorls often appearing white ing of small rheocrenes (16-27"C, 2000- 10, duc to erosion. Aperture broadly ovate, 150 micromhoslcm), snails were common apertural plane near-parallcllO coiling axis on a ll substrates. (Fig. 15b). In ner li p thickened, reflected Etymology. - Re ferring to d is tribution above, adnate to small ponion of or slightly along Amargosa River. separated from body whorl. Outer li p Remarks. - Distinguished from p, micro­ straight. thin. Umbilicus chink-like to coccus by globose shell; large penis; large broadly open. Spiral striations sometimes penial fila ment; small, narrow penial lobe; prominent on tcleoconch. and highly reduced glandular ridge. Ranges Dark, brown-black epi thelial pigment on of these very simi lar snails (previously con­ most of snout (except distal tip), al ong an­ fused by Hcrshle r and Sada 1987:788) are terior and posterior edges of " neck" (ccntral contiguous. area lighter), on part or all of operculigcrous Pyrgulopsis micrococcus (Pilsbry, 1893) lobe. Tentacles lightly pigmented. Oasis Valley springsnail Radular (Fig. 16) form ul a: 5(6)-1-5(6)/ Figs. 17c, d, 20-25 1-1 ,3-1-4,27-28,24-26 (from para types). Central tooth broadly trapezoidal: basal Amnicola micrococclls Pilsbry in Steams, process moderately excavated. Penis (Figs. 1893:277. VOLUME 102. NU MBER I 183

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Fig. 1. Sampling localities in Death Valley and Amargosa Ba~in.

Malerial examined.-Nevad a: Nye CO.; 857966, 21 Mar 1987.-Lower spring in Spring in Oasis Valley (204), USNM Johnson Canyon (220), -853509, 18 Feb -850297. 18 Nov 1985; 857961, 16 Mar 1985; 857967, 21 Mar 1987.-Stream in 1987.-CaneSpring (239), -857936, 10]uly Jail Canyon (196), -8535 13,14 Fcb 1985; 1986.-Califomia: InyoCo.; Northernmost 857970, 19 Mar I 987.-Strcam in Surprise of Tecopa Hot Springs (2 10), -853502, 12- Canyon (194). -857937, 19 Mar 1987; 13 Mar 1985; 857962, 16 Mar 1987.­ Stream in Pleasant Canyon (190), - 857940. Shoshone Spring (211), -853501, 12 Mar 19 Mar 1987. - China Garden Spri ng (184), 1985 ; 857962, 16 Mar 1987.-Grapcvine 853511. 9 Feb 1985.-Stream below Dar­ Springs (241): northwest-most spring in win Falls (184). 857969, 23 Mar 1987.­ complex, 853503, I Mar 1985; Spring Spring above Darwin Falls (184). -85351 2. emerging below limestone bench northwest 10 Feb 1985.-Stream in Snow Canyon of ranch, 853504, 26 Feb 1985; Spring on (182). - 857938. 16 Sep 1987. J . Aardahl!­ bench above ranch, 853505, 25 Feb 1985; Stream in Knight Canyon (180). - 857939, Spring al ranch, -853506, 25 Feb 1985: 14 Apr 1981.-Tennessee Spring (178), 857964, 14 Mar 1985.-Spring 1.6 km east - 853514, 4 Mar 1985; 857971, 13 May of SCO tty's Castle (2 42), -853507. 26 Feb 1987. -Saline Marsh (152), -853510. 10 Feb 1985; 857965, 14 Mar 1987.-Hanaupah 1985; 857968, 28 Mar 1987. Spring (222), -8 53508, 16 Feb 1985: Diagnosis. -A small·sized species, with '" PROCEEDINGS OF THE BIO LOGICA L SOCiETY O F WASHINGTON

Fig. 8. Photographs of seleded springsnai1 localities: 3. stream in Couonball Marsh. Death Valley. I)-pe locaht} of T. Mllmo (3·10·87; photo by P. Rowlands): h. Grapevine Springs. Death VaUey_ type localily of T. margae and T 'O~ londs; is on lhe laTie 1ra,-cn;nc mound (3-1 4-87); c. Saline Marsh. Saline Valle) (3-29-81); d. Sand Canyon stream. Indian Wells Valle)' (3-26·87): c, Fish Slou&h. O"'"Cns Valley. spring sou~ (including type localily of P. penll,balaj ' -;sible in 10""'" ponion of photograph (ca. 1975; aerial photo by P. Piner): f. "NE" Spring a1 fish Slough (2-7-85).

globose to Ovate-conic shelL Penis with bulging of whorls. Body whorl 65-89% of moderate-sized lobe; distal edge oflobc usu­ shell hcight. Aperture ovate, a pcrtural plane ally ornamented with glandular ridge. near-parallel 10 coiling axis (Fig. 2 Ib). In ner DeSCrIption. -Shell (Figs. 20, 21) 1.1-3.1 lip moderatcly thickened and reflectcd. usu­ mm high, height/width. 99-190%. Whorls. ally slightly separated from body whorl. 3.25-5.0. well-rounded, with slight shoul­ Outer lip straight, thin. Umbilicus chink­ ders. Spire often irregularly convex due to li ke to moderately open. VOLU ME 102. NU MBER I '"

Fig.. 9. Photograph, of springsnaillocalilies: 3, Wann Springs. O",-ens Vaney (4·19·87): b, Small spring S of Wa rren Lake. O ..-ens Valley (4.18_87): c. Spring in Owens Gorge. O",-ens Valle)' (5-7-87): d. H OI Creek. LoTlJ Valle), (4_23·87). '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. SEM micrographs of P. aardahll from spring 1\ Bramlette Ranch: a. Hololypc. USNM 860406 (shell height. 3.26 mm): b-d, Paralypcs, USNM 857951 (scale of"b" and "e" as in "a": bar - ISO I'm). VOLUME 102, NUMBER I '" Head/foot epithelial piament ranging from absent to near-uniformly black (lighter on central ponions of sides of '·neck"). Radular (FilS. 22, 23) formula: (4-7)-1- (4-7)/1-1 , (2-4)-1-{3-6), 17-2b, 1 ~-32 (from numerous populations). Central tooth moderately to broadly trapezoidal; basal process moderately excavated. Penis (FilS. 17c. d, 24b-i) barely extend­ ing beyond mantle collar, moderately thick­ ened, longer than wide. Filament slender, FiJ, II. Riehl IateTal vicw ~oot of P . (I(IT· do.hli. USNM 85795 I. sprin,at Bram1clle Ranch .Sup­ usually extending we ll beyond edge of lobe. p .... indicate pi,menled an:as. Lobe hiahly variable in size and shape. ta­ pering slightly distally. Glandular ridge (rarely absent) small, positioned on or near (97), -857950, 26 Apr 1987.-Warm Springs distal edge of lobe. Capsule gland small er (98), Main spring, -853544, 7 Feb 1985, than albumen gland (Fig. 24a). Seminal re­ 857988, 19 Apr 1987; Small spring north ceptacle small, positioned la rgely anterior ofahove, l'l5J545, 7 ."c b 1 9~5; ~57~l'l~, I ~ to bursa copulatrix. Bun&. pear-shaped, Apr 1987. Mono Co.: Stream in canyon S small to fairl y large relalive to pallial ovi­ of Piute Creek (104), 860404 (holotype), duct, most oflength positioned on albumen ·857955 (paratypes), 8 May 1987.-Springs gland. on bench south of Piute Creek (103), North Type locality. -Small spring in Oasis Val­ spring, 857986, 2 1 Apr 1987; South spring, ley, Nyc County, Nevada (precise location 857987, 22 Apr 1987. unknown). Diagnosis. -A small- to moderatc-sized Distribution and habilat.-A widespread species with globose to ovatc-conic shell. form (contrary to Taylor 1985:317) found Pcnis large relati vc to hcad/foot, filament in smaJl springs and stream outflows (see short, lobe enlarged, ventral swelling pres­ Figs. 8b, c) of Amargosa Ri ver drainage as ent. Penial glandular ridges, 2-6. well as interior drainage in Death, Pana­ Descriprion. -ShelJ (Figs. 26a~, 27, 28) mint, and Saline Valleys (5-32OC, 300-4000 1.5_2.8 mm hiih, height/width, 110-160%. micromhoslcm) (Fig. 25). Typically com­ Whorls, 3.0-4.25, wcll-roundcd, sutures mon on stone, bits oftravenine, watercress, slightly impmsed. Protoconch sometimes and plant debri s. slightly tilted. Body whorl in flated. height 74-90% or that of the shell. Apical whorl s Pyrgu/opsis o .....ensensis. new species sometimes eroded, shell often covcred by Owens Valley springsnail deposits. Aperturc ovate. angled abovc; Figs. 26a~, 2'-32 moderate to large in size. Aperture plane Material examined. -Cali fornia: In yo near-parallel to somewhal tiltcd relative to Co.; Spring ca. 1.6 km south of Mule Spri ng coiling axis (Figs. 27b, 28b). Inncr lip slight­ (92), USNM 85354 1, 5 Dec 1987, D. G iu­ ly to considerably th ickened and re ncctcd. liani.-Mule Spri ng (93), -853540,19 Nov adnate 10 or (more commonly) sligh tly sep­ 1985, D. Wong; 857983, 20 Apr 1987. ­ arated from body whorl. Outer lip thin. Spri ng at Graham Ranch (9 4), -853543, 8 straight or slightly sinuate. Umbilicus chink­ I-'eb 19~ 5; ~57~~5 , l~ Apr 19S7.-Wilk­ like to modcrately open. erson Springs (96), -853542. 8 Feb 1985: Dark brown or grey-black epi thelial pig­ 857984, 18 Apr 1987 .-Spring at Toll House ment on snout, peripheral portions or '88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 12. Radula of P aardahli. USNM 85195 I, spring at Bramleuc Ranch.: a. Ccn1rals (bar - 11 I'm): b, Laterals (bar - 8.6 I'm): c. Inner marginal (bar - 6.7 "m); d. OUlcr marginals (har " 6.0 I'm).

"ncck," and dorsal surfaces of tentacles. Radular (Figs. 29, 30) formula: 5(6}-1- Dark subepithelial granules prominent in 5(6)11-1, 3-1-3(4), 18-23, 25-37 (from central areas of sides and sometimes along three populations). Centra! tooth broadly posterior edges of"ncck." trapezoidal; basal process well-excavalcd. VOLUME 102. NUMBER I '" Penis (Fig. 3 1~i ) ex tending well beyond mantle collar, elongate. Narrow fi lament 'v-~_," shorter than lobe. Lobe massive, tapering only slightly, distaJ edge blu nt. Edge orlobe ornamented with single elongate or two shorter glandular ridges. Dorsal penis some­ o times having small gland ul ar ridges on lobe and/or near base of filament. Ventral swell­ ing small, located filam ent base. Ridge or­ namenting swelling variably-sized. Fila­ \ ment darkly pigmented ....' th sub-epithelial granules along most of length (Fig. 3Ib). Capsule gland equal to (Fig. 31 a) or slightly smaller than albumen gland . Seminal re­ ceptacle minute, positioned largely or en­ tirel y anterior to bursa. Bursa pear-shaped. small- to moderate-sized. wi lh about half of len&th posteri or to albumen eland . Type locality - Stream in canyon S of Piute Creek, Mono Co., California. Distribution and habitat.- Found along Fig.. 13. Reproductive anatomy of P. aardahli. escarpments of White and Inyo Mountains USNM 857951. spring at Bramlett~ Ranch: a, Left cast side ofQ....'ens Valley (Fig. 32). Habitat lateral view ofpaltial oviduct comple,,:!>-e. Penes (b. consisting of small springbrooks (I 0-29"C, c. do~l: d. ~. '·em",I). Suppled areaS on penes Ind,ca!e glandular ridges (e ~closed stipples) and pigmented re­ 300-780 micromhoslcm) (Fig. 9a) where gions (open). Ag - albumen gland; Bu - hu~ copu­ snails typically common in watercress and! latri,,: Cg - capSule gland: Cga - capsule gland open­ or on bits of travertine and stone. Syntopic Ing: Emc - poSteno.-end ofpallial ca\'tt)~ 0" - o,iduct: with P. wong; in spring at Toll House. PC - penial filament: Plo - penial lobe: Sr - ~mlnal Erymolog)'- Rcferri ng 10 occurrence in receptacle: Vc - wntral channel of capSule gland. Q....~ns Valley. Remarks. - Distinguished from similar P. Diagnosis. - A large-sized species with penurbata by shell appearance. more en­ thickened, low-conical shell. Penis large, larged penial lobe. and (usually) more nu­ lobe and filament small, ventral swelling merous penial glandular ridges. present. Penial glandular ridges. 3 or 4. Description. -Shell (Figs. 26e-g. 33) 2.7- 4.0 mm high. height/width. 130-180%. p}'rgulopsis penurbala, new spcries Whorls, 4.25-5.0, only slightly convex ...... ith Fish Slough springsnail pronounced sub-sutural angulations. Body Figs. 26c--g, 33---36 whorl inflated. 71 - 85% of shell height. Ap­ Material examined.-Califom ia: Mono enure broadly ovate. Apertural plane tilted Co.: Fish Slough: "Northwest Springs:' relative 10 coiling axis (Fig. 33b). Inner lip South spring (76), USNM 860407 (holo­ usually well-thickened and reflected. broad­ type), - 853546 (paratypes). 7 Feb 1985: ly adnale to body whorl. OUler lip slraight. 857990,24 Apr 1987: "Nonhwcst Springs," thin or moderately thickened. Umbilicus Non h spri ne: (76). 853541. 7 Feb 1985: chi nk-like. 857991. 20 Apr 1987: "NE Spring" (77). Dark brown-black epithelial pigmenl -853548.7 Feb 1985: 857992, 20 Apr 1987: prominent on snout. tentacles. and periph­ " BlM Spring" (75). -853549. 7 Feb 1985. eral ponions of sides of head/fool. Pigment .90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

""

~ -, ,,

•

-'­, , o " "" " " .- "" u" N/ ,,-, , b ..... ~J " • " " .. KB"m.'.,. " 8' 11 7'

fig. 14. Map showing dimibution of P. aQrdahli. either light or absent on central portions of thin, considerably longer than wide. Fila­ sides of head/fool. ment both narrow and short, barely extend­ Radular (Fig. 34) formula: 5-1-51 I-I, 3- ing beyond lip of lobe. Lobe tapcnng only 1-4(5), 18-19, 29- 31 (from para\ypcs). slightly distally, bearing elongate glandular Central tooth broadly trapezoidal, basal ridge along distal edge. Dorsal surface of process well-excavated. Penis (Fig. 35b-e) penis with small ridge proximal to base of extending well beyond edge of mantle collar, fIIamenL Ventral swelling arising slightly VOLUME 102. NUMBER 1 '"

Fig. 15. SE M micrQgraphs of P. amargosae From Saratoga Spring: a. Holot)'pe. USNM 860401 (2.18 mm): b. c (bar - IJ6 I'm). Paratypcs. USNM 853515 '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 16. Radula of P. amargOSQi', USNM 853515, Sar.uoga Spring: a, Centrals (bar - 12J'm); b, LarenoJs (bar - 8.6 ~m): c. Inner marginals (bar - 8.6 I'm): d. Oulcr marginal (bar - 3.8 I'm). proximal 10 base of fIlament; ridge (rarely 10 albumen gland (Fig. 35a). Bursa small , two) on swelling elongate. Filament dark­ with ca. 50% oflcnglh posterior to albumen ened along much of length by sub·cpithclial gland. Seminal reccplaclc mi nute, posi­ pigment granules. Capsule gland sub-equal tioned entirely anterior to bursa. VOLUME 102. NUMBER I '93

Fi&. 17. SEM micrOVolptis ofh~ad·foot and dorsal penes of P. Qmtlrg051U and P. mic; oroccus. a (bar " 0.43 mm). b (bar - 0.38 mm). P. mna;losa<" USN M 853515, Saratoga Sprin8: c (bar - 0.3 mm). d (bar - 200 I'm). P. micrococcus. USN M 857961. spring in Oa$i$ Valley. '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tppp lo('ality_ - "Norlhwest Springs," south springpool. Fish Slough, Mono Co., California. Distribution and habitat. - Found in three of the four main springs in Fish Slough (ex­ tinct in "BLM Spring") (Figs. 8e, f, 36). Springs iss ui ng relatively cool (22-26"C), soft (200-360 micromhos/cm) wa ter. Snails found only in small vestiges of rheocrcne habitat at various small orifices in north­ west springs (rare al both sites on vegetation and stones), and at beginning ofoul How from "NE Spring" (common in Chara). Etymo!ogy. - Referring to various distur­ bances to endemic locale, including severe reduction of original rhcocrcnc habitat as unanticipated side effect of efforts to con­ serve Owens pupfish (Miller and Pister 1971).

Pyrgulopsis cf. slearnsiana Fig. 18. Reproduccive anatomy of P. cmargoS

Fig. 19. Map showing d;~tribu'ion of P. amargosai'. projecting distal to base of fi lament. Tip of Filament with dark sub-epithelial pigment lobe often ornamented with small glandular streak (Fig. 39b). Albumen gland sub-equal ridge; similar, single ridges on dorsal penial \0 capsule gland (Fig. 39a). Seminal recep­ surface about halfway from base of penis to tacle small, positioned anterior \0 bursa base of fi lament, on ventral surface near copulalrix. Bursa copulMri" small relMive (sometimes on) inner edge. The latter ridge 10 capsule gland, positioned panly posterior enlarged, borne on low swelling (Fig. 3ge). 10 gland. ". PROCEEDINGS OF THE BIOLOGICAL SOCIETY O F WASHI NGTON

h

Fig. 20. Camera lucida drawings of shells of P. ",ic~ a, USNM 857936. Cane Sprinl (2.2' mm); b. USNM 853501. Shoshone Spring (1.96 mm): C, USNM 853502, Tecopa HOI Springs (1.9() mm); d. USNM 853506. Grapevine Springs (2.16 mm); c, USNM 853508. Hanaupah Spring (2.26 mm); f. USNM 8SJSIO. Saline Mal"Sh (1.96 mm); s- USNM 853'12. ,o.~n Fall. (1 .94 mm); h. USNM 857931, S" rpri ... Canyon sl«:am (l .SS mm).

Type locality. - Near Oakland, Alameda identi ty of the snails from Indian Wells Val­ County. California. ley. Distribution and habitat. - Distribution given by Taylor (1981: 152) as "Central Cal­ Pyrgulopsis wongi. new species ifornia, from Sonoma County to Monterey Wong's springsnail County along the coast, and inland in the Figs. 41-47 foo lhills of the Sierra Nevada." Found in the study area in twO moderalc-sizcd, cool Material examined. -California: fnyo. streams (16--2O"C, 550 micromhoslcm) sep­ Co.; Spring at Little Lake, cast of HW 395 arated by 25 km along eastern slope of Sier­ (10), USNM ·853521, 9 Feb 1985.-HOS­ ra Nevada in Indian Wells VaHey (Fig. 40). back Creek (Monachc Min. quad.) (20), Snails common in watercress and Chara. ·853522,28 Jun 1985, D. Wong.-Sum mit Remarks. -Shell s of the Indian Wells Creek (21), 853523, 12 Apr 1987.- Spring Valley snails closely resemble P. slearnsiana ca. 3.0 k.m north of Summit Creek (2 1), from the San Francisco Bay area and their ·853524, 12 Apr 1987.-Spring at Cabin penes conform to a general description for Bar Ranch (25), 853525, 12 Oct 1987, J. the species provided by Gregg and Taylor GOldberg and M. Shumway.-Lubkin Creek (1965: I 07). Additional study. including ex­ (34), 853526, 16 Apr 1987 .-Spring feed ing amination of anatomy of topotypes or near­ Lubkin Creek (34), 857942, 25 Apr 1987.­ topotypes, will be necessary 10 confirm Spring along east side of Tuttlc Creek (35), VOLUME 102, NUMBER 1 '"

Fi&. 21. SE M m.crogn,phsof P. microco«us: a.. USNM 853502. Trcopa HOI Spnngs (1.98 mm): b. USNM 853512. sprin& aoo"e Da rwin Fal:s: c. USN M 8j)j 12. Te<:opa HOI Spnngs (bar - 120 ~ m). ". PROCEEDINGS OF THE BI OLOGICAL SOCIETY OF WASHI NGTON

Fig. 22. Central radular teeth or P. mic"xoccus: a, USNM 857936. Cane Spring (bar - 1.5 I'm): b. USN M 857963. Tecopa Hot Sprinll$ (bar - 7.5 I'm ): C. USNM 857965. spring E ofScolly's C""tle (bar - 7.5S'm); d. USN M 85 7968. Saline Marsh (bar - 8.6 I'm).

857943, 30 Apr 1987.-Hogback Creek Gi ul iani. -Spring on hill soulh of Warren (Lone Pine quad.)(36), "853527, 8 Feb 1985; La ke (57), *853530, 18 Apr 1987.-Spri ng 857976, 16 Apr 1987.-Boron Springs (40), in canyon south of Shannon Canyon (59), *853528,17 Apr 1987. - Stream in Charlie 857944,20 Apr 1987.-Spring along north Canyon (43), *853529, 12 J UII 1987, D. side of upper Pine Creek (69), *853531 , 24 VOLUME 102, NUMBER I '"

Fig. 23. Lateral and marJinal.adular teclh of P. micrococcw: a. Lateral., USNM 857936. Canc Sprin8 (bar - 6.0 lIm): b. Laterals. USNM 857966, Hanaupah Spring (bar - 7.5 lIm): C, [nncr marJinals. USNM 857937. Surprise Canyon stream (bar - 5.0 lIm): d. Outer marJinals, USNM 857963. Tecopa Hot Springs (bar - 4.3 lIm).

Apr 1987; 857977, 15 Scp 1987, D. Wong.­ 857945, 3 May 1987.-Spring in Owens West spring in Birchim Canyon (70), 860403 Gorge (74), ·853532, 14 May 1985. D. (holotype), ·857941 (paratypes), 3 May Wong; 857978, 8 May 1987. - French Spring 1987. - East spring in Birch im Canyon (70), (89), · 857947.4 May 1987.-Barrel Springs 200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Description. -Shell (Figs. 41 , 42) 1.2-3.0 mm high, heighVwidth, 100-160%. Whorls, 3.25-4.5, well rounded, with slightly an­ gulated shoulders. Body whorl 76-87% of shell height. Aperture broadly ovate, angled above. Apcnural plane slightly tilted rela­ tive to coiling axis (Fig. 42c). Inner lip slightly thickened, usually only minimally reHeeted, either adnale 10 portion of or slightly separated from body whorl. Outer lip straight, thin. Umbilicus chink-like to moderately open. Dark grey-black epithelial pigment on snout, proximal tcnlaclcs, and peripheral ponions of sides of head/ foot. Entirety of head/foot often very dark, Central portions of sides of head/ fOOl usuall y wi th densely clustered dark, subepithelial pigmenl gran­ ules. Radular (Figs. 43, 44) formula: (4-7)-1- (4-7)/1-1 , (2-5}-1-(3-6), 17-26,20-34 (from numerous populations). Central tooth FiJ. 24. RCPfoouctive anatomy of P. miCTOCOCCW: a, left lateral view of pallial oviduct complcx. USNM relatively square-shaped; basal process well ­ 857961 , spling in Oasis Valley; 1:>-;. Penes (aU dorsal excavated. Penis (Figs. 45b-1, 46) flattened, excep. "i")(b. i. USNM 857936, CaneSpring;c, USNM longer than wide. Filament fnirl > broad, 853505, Gupevinc Springs; d. USNM 857962. Shosh­ short relative to le ngth of remaining penis. one Spring: e, USNM 857963, ncopa HOI Springs; f, Lobe slightly shorter than filament, broad, USNM 851967. Johnson Canyon spring: g, USNM often widening distally. Glandular ridge ar­ 857971 . T c~ncsscc Spring: h. USNM 853512. spring above Da",in Falls). rangement typically as follows: I) elongate ridge li ning edge of lobe; 2) similar, stalked ridge on ventral surface just pro)!.imal to (9 1), *857948, 18 Aug 198 7. ~Spring at Toll lobe; 3) sim ilar ridge (rarely very reduced) House (97), *857949, 26 Apr 1987. ~CoTTa l on fleshy projection of ventral surface pro)!.­ Springs (160), *853535, 8 Feb 1985; 857979, imal to above; 4) elongate ridge covering 29 Mar 1987.-Antelope Spring (1 62), proximal-most two-thirds of dor>al fila­ *853536,8 Feb 1985. Mono Co. ; Spring ment; 5) very elongate ridge on dorsal sur­ along Marble Creek (82), *853533, 22 Apr face curving from just pro)!.imal to filament 1987. -Springs at north end or Blind Spring to lobe; 6) shorter ridges on dor>al surface Valley (86); North Spring, +8 57946, 21 Apr along right side of (5) and in area between 1987; South spring, 853534, 21 Apr 1987.­ base of filam ent and edge of lobe. Modifi­ Layton Springs (12 2), *8 53537, 6 Feb 198 5; cations of above include addition of a few 857980, 23 Apr 1987. - River Springs (132), small ridges on central dorsal surrace, ad­ *853538,6 Feb 1985; 85798 1, 21 Apr 1987. dition or a si ngle small ridge (sometimes Diagn05is. -A small- to moderate-sized stalked) on ventral surface, apparent rrag­ species with globose to low conical shell. mentation of ridges (particularly" I," "5"); Penis massive, with large filament and mod­ apparent merging ofadjacent ridges on dor­ erate-sized lobe. Ventral penis wi th two sal surface. Filament darkened with sub-e p­ prominent swellings bearing glandular ithclial pigment along most of length. Cap­ ridges. Glandular ridges, 7-12. sule gland sub-equal to albumen gland (Fig. VOLUM E 102. NUM BER I 20' , , , ,~,-

'>c'"':.:. f' l- (""'~"'() u-? ::-:! , <;:fJ !;;:;~ i'" ""'", "- "- , •" _0 -~ , ,,' ,, ,<' , , ~~, , , '" ,, , , ~ , "'~\ , , 0, ., , • , -J •• ::;:, ,. , • • · , , • -• , • • , , •y "' . , ASh Meadows " , , • \-/"', • , ~1 , I L , , , , . y , , " , <. " , 0 • "I~ y , • • " , , • •• 1'" ". - • "c'". •• '. " N! "• ,,', , tv- 'J '0 • " " " .., ~ llo ... ' ... '. I ' '8' I I U'

J",.. 25. Map showing distribution of P. mlCro<:C<:cUS_ Snail occurs at 15 localities in Ash Meado""s.

45a). Sem inal receptacle small, positioned lillie Lake, and along western side of val­ lateral to anterior bursa copulatrill. Bursa ley from French Spring to Marble Creek, pear-shaped, massive, wi th about one-third Also found in a few sites in Long. Adobe, of length posterior to albumen gland. and Deep Spri ngs Valleys (Fig, 47). Habitat Type locality. -Spring in Birchim Can­ includes seeps and spring-fed streams of yon north of Bishop, lnyo Co" California. small-moderate size (9.5-22°C, 85-450 Distribution and habitat. - Widespread in micromhoslcm) (Fig, 9b. c). Snails typically Owens Vall ey along eastern escarpmen t of common in watercress and/or on small bits Sierra Nevada from Pine Creek south to of travertine and stone. 202 PROCEEDINGS OF THE BIO lOGICAL SOCIETY OF WASH INGTON

g

Fig. 26. Camera luciaa (Iraw;ngs Of shells or 1'. Olnurbala: a-d. 1'. O"WISl''lSO (a. USNM 853540. Mule Spring [2.! mmJ: b. USNM 857950. spring al Toll House [2.3 mmJ: c. USNM 8H9SS, SU'eam in canyon S of Pi ute Creek p.08 mrn1: d. USNM 853543, spring al Graham Ranch [2 .16 mm]): ~ F.perturbaui (e. USNM 853546. Fish Slough. "Nonhwl"St Springs:- [3.93 mm): f. USNM 853549, Fish Slough, " BLM Sprina," 13.22 mm!: g. USNM 853548. f ish Slough. "NE S»rins" [3.97 mm)).

Etymology. - Named after Darrell Wong, spring on limestone bench, USNM 860408 California Fish and Game, for his assistance (holotype), ·857952 (paratypcs), 14 Mar during the project and interest in conserving \987; Spring on li mestone bench above Owens Valley springsnails. ranch 853555, 25 Feb 1987; 857996, 14 Remarks. - Simila r in penial morphology Mar 1987. to P. caiiforniensis (Gregg & Taylor) from Diagnosis. - A small -sized species with western California (Taylor 1981: 154). Dis­ narrow. turri form shell. Penis small. with 3 ti nguis hed fro m above by globose shell. and papillae on inner curvature, and a single more numerous stalked glandul ar ri dges on papilla (sometimes abse nt) on (J uter cur­ ventral penial surface. vature. Distal tip of penis with elongate Genus Tryonia Stimpson, 1865 swelling on inner side. Trronia margal?, new species Description.-Shell (Fig. 48) U-2.9 mm Grapevine Springs elongate tryonia high, height/width, 250-300%. Whorls, 4.5- Figs. 48-50, 5 1a , 52 6.25, rounded, wilh angulated shoulders. J faterialexamined. -California: Inyo Sutures often deeply impressed. Apex bl unt. Co.; Grapevine Springs (24 1); (upper) Warm protoconch slightly depressed (Fig. 48g). VOLUME 102, NUMBER 1 203

Fit,. 27. SEM mIcrographs of P. ol<"ensi'/lSis from SIT~ am in canyon S of Pi ute Creek: a. HolotyP<', USNM 860404 (2.0 mm): b-d. Paratypcs. USNM 851955 (bars - 120 I'm, ! 76 I

Body whorl 45-55% of shell height. Marl tc1coconch. often elevated al irregular in­ frequently covering apical whorls, with near­ teryals. Spiral sculpture consisting of weak entirety of shell sometimes blanketed. Ap­ striati ons. erture narrowly ovate. Inner lip well-reflect­ Dark brown- black epithelial pigment ed, slightly thickened, adnale 10 or slightly covering most ofhcadlfoot. although some­ separated from body whorl. Outer lip si n­ ti mes absent on tentacles. and lighter on uate. thin (Fie. 48d. O. Umbilicus ehink­ snout and central pon ion of sides of ' 'neck ... li ke. Protoconch near-smooth (Fig. 48g). Radular ( Fi g~. 49, 50) formula: 4(5}-I- CollabraI growth lines we ll developed on 4(5 )/ 1(2}-1(2), 2(3}-1-3(4). 20-21. 22-28. 2(), PROCEEDINGS OF THE BIOLOGICAL S

Fij. 28. SEM microwaphs of P. o"'enseltsis from Warm Springs: a (1.92 mm), b, d, USNM 8SJS44. main spring; c, USNM 853545. spring N of above (bar - 136 "m).

Central tooth broadly trapezoidal, basal on inner edge. Brown-black cpilhe li al pig­ cusps large, basal process well excavated. ment usua ll y prominent near distal tip, al Central cusps of lateral tooth only slightly bases of two distal papillae. Lighter pigment enlarged (Fig. 49c, d). Penis (Fig. 51a) ex­ dusting much of dorsal surfaces of penis and tending only sl ightl y beyond edge of mantle 2 proximal papillae. collar. Enlarged papilla located on outer edge Type localily. -Warm spring on lime­ of penis al base. Two smaller papillae po­ stone bench in Grapevine Springs complex, sitioned along distal half on inner edge; sim­ nonhem Death Valley, Inyo Co., Califor­ ilar small papilla usually found near base nia. VOLUME 102, NUMBER 1 20S

Fig. 29. Radula of P. owensl'ns;', USNM 857955, stream in canyon S ofPiL1le Creek: a, Centrals (bar - 10 $Om): b.l..aterals (bar - 6.7 I'm): C. Inner marginal (bar - 6.0 I'm): d. Outer ",arginals (bar - 6.0 I'm).

Distribution and habitat. - Found in two mineralized ( 1050-1075 micromhos/cm), sites (separated by about I km) among but d iffered considerably in temperature Grapevine Springs in northern Death Val­ (west spring, 26-27"C; east, 36-37"C). Syn­ ley (Fig. 52). Habitat consists of upper por­ topic with T. rOll"landsi in the thermal spring. tions of shallow spri ne brooks Howine over Elymn!ngy.-From Latin marga. mean­ travertine (Fig. 8b), on which snails were ing marl , and referring to occurrence of snail moderately common. Sprin~s were highly in travertine springs. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 30. Radula of P. Ok·emensis. USNM 857988. Wann Springs (main spri"il: 3. Cenlral5 (bar - 6.7 pm); b. Lalerals (bar _ 6.0 pm); c. Inner marginal (bar - 6.0 pm); d. Ouler marginals (bar - 3.8 pm). VOLUME 102, NUMBER 1

Tryonia pro/ea (Gould, 1855) b Desert tryonia Figs. 52- 54 A m nicola prou'a Gould t 855: 129. Melania exigua Conrad 1855:269. Pyrgulopsis blakeana Taylor 1950:30. , Pyrgulopsis cahuillarnm Ta}lor 1950:3 1. o Material examined. -California: Mono Co.; Hot Creek, USNM '"857954, 23 Apr 1987. Diagnosis. - A moderate to large-sized species (shell heighl typically 3-7 mm) with elongate-conic to turrifo rm shell. Shell smooth or with spiral and/or col1abral d , sculpture varying from faint striations to elevated ridges. Description of H OI Creek popllfminn._ Shell (Fig. 53) 3.0-4.5 mm high, height/ wid th, 2 10-242%. Whorls, 4.75-6.25, rounded. Sutures moderately impressed. ; Body whorl 55-61% of shell height. Pro­ toconch slighll y depressed (Fig. 53b), apex ofaduits invariably highly eroded . Aperture ovate, apertural plane tilted re lative to coil­ ing axis. Inner lip only sligh tly thickened and reHected, broadly adnate to body whorl. Fig.31. Reproduceive anaeomy of P. o"-enscnsis: Outer li p th in, sligh tl y sinuate. Umbilicus a. Len laecral view ofpallia] oviduce complex. USNM chink-like, near-absent. Smooth surface of 857955. scream in canyon S of Piuee Creek: b-i. Penes (b [dorsal). i [vcnerall. USNM 857955. scrcam in can­ protoconch lined wi th well -spaced rows of yon S of Piuee Crttk: c [dorsall. f [vcnerall. USNM numerous, shall ow perforations (Fig. 53b, 857983. Mule Spring; d [dorsall. g [vcnuall. h [veneral). c). Teleoconch wit h strong growth lines; 10- USNM 857988. Warm Spring (main spring): c [ven­ 20 spiral li nes prominent on mi ddle whorls, Ira]]. USNM 857989. warm Springs [spring N of sometimes extend ing to apen ure. above]). "b" ehrough "i" drawn co samc scale. Brown epithelial pigment dense on most of snout. Black patches of subepithelial pig­ ment along edges of sides of head/foot. Pal­ bly in Riverside County, California [based lial roof with paired, black pIgment patches on Blake 1857)). extending along length of ctenidium. Distribution alld habitat. - "Western Utah Radular (Fig. 54) formula: 5(6}-l-5(6)! to southeastern Cali fornia, adjacent Baja 2(3)-2(3), (3- 5)-1-(6-8), 35-36, 28-31 California , and southeastern Arizona" (from single po pulation). Central tooth (Taylor 1981: I 55), with only two living broadly trapezoidal; basal cusps large, basal populations known from California: Ihat of process moderately excavated. No males Hot Creek (Fi g. 52) and Dos Palmas Spri ng fo und among the more than 100 specimens in Riverside COIJnly (Taylor 198 1: 154). AI collected . Hot Creek (Fig. 9d), snails were rare on var­ Type locality.-Colorado Desert (proba- ious submergent macrophytes lining sides 208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WAS HINGTON

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• • •• " J , • " , , d "'2"

" "" " C'"" NI <, b

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Fig. 32. Map showing dislribution of P. owens"lts;s. of the swi ftly flowing thermal stream (32"<:, by analO mical comparison with snails from 480 micromhoslcm). Dos Palmas Spring (thaI re present near to­ Remarks. - The taxonomic concept of this potypes) would be desirable. form is vague (Taylor 1966:53-54) and sta­ tus of the few living (and mosily widely sep­ Tryonia robus/a, new species Robust lryonia arated) populations (Taylor 1985:fig. 35) rc~ mains unclear. T he Hot Creek populati on Figs. Sid, 52, 55-58 is referable \0 T prOiea on basis of shell Material examined. -California: In yo form, size, and sculpture, but confi rmation Co.; Nevares Springs (230); Spring on trav- VOLUME 102, NUMBER 1 '"

Fia_ 33. SEM microg.rnphs of P. fH-'rlurbata rrom Fish Slough. ··Nonh ..-e~t Sprinp:' south Wring: a. b. Holotype. USNM 860407 (3.57 mm): c. Parntype. USNM 853456 (bar - 200 I'm). crtine mound, USNM 860411 (holotype). 1985-Travcrtine Springs: South spring. -853557 (paratypes), 2 Feb 1985; 857999, -853559.2 Feb 1985: 857959.12 Mar 1987: 16 Nov 1985; 857958, 12 Mar 1987; Spring North spring, 857960.12 Mar 1987. issuing from base of mound. 853558.2 Feb Diagnosis.-A small-sized species with '" PROCEEDINGS OF THE BIOlOGICAL SOCIETY OF WASH INGTON

Fig. 34. Radula of P. per/lirbata. USNM 857990, Fish Slough, "NonhweSISprings," SOUlh spring:a. Centrals (bar - 13 .~ I'm); b. Laterals (bar - 10 I'm); C, Lalerals and inner marginals (bar - 12 I'm): d, Oute, marginaJs (bar - 10 ~m ). unusually broadly conical shell for genus. high, height/width, 140-170%. Whorls, Penis moderate-sized, with single papill a on 3.75-4.75, rounded , with pronounced outer curvature. shoulders. Sutures impressed. Body whorl Description. _Shell (Fig. 55) L 1_2. 1 mm 68-80% of shell heiellt Protocon~h (Fig"'_ VOLUME 102, NUMBER I '"

56a-c) simple, sometimes eroded. Aperture b near-ovate, sometimes sharply angled above; apertural plane slightly tilted relative to coiling axis. Inner lip thickened, mod­ erately reHected, usually slightly separated o from body whorl. Outer lip thin, ncar­ straight. Umbilicus chink-like. Protoconch surface wrinkled (Figs. 56a-c). Teleoconch often with weak spiral striations, particu­ larly on upper whorls. Operculum (Fig. 56d) thin, paucispiral, with 2.5-3.0 rapidly expanding whorls. Brown-black melanic pigment dark on d , snout and posterior edge of"neck," variable on central portions of sides of head/foot; sub-epithelial pigment granules prominent in latter. Mantle roofepitheJial pigment Iyp­ ically unifo rmly dark brown_black. Radular (Fig. 57) formula: (5-7)-1-(5-7)/ •

1-1 , 3(4)-1-5(6),24,22-27 (from two pop­ Fig. 35. Reproductive anatomy of P. penurbFuneral Mountains in mid·eastern Death spring on limestone bench, USNM 860409 Valley (Fig. 52). Snails found commonly on (holotype), *857953 (paratypes), 14 Mar plant debris and travertine in shallow out­ 1987; (lower) Warm spring on limestone flows offour thermal springs (30-37"C, 900- bench, 857997, 14 Mar 1987. 975 micromhos/cm). Extinct at Texas Diagnosis. - A small·sized species wi th Sprinp (located between above complexes). elongate-conic shell. Penis large, wi th single Etymology. - Referring to unusually papilla on outside curvature and 2 papillae broad shell for genus. on inside curvature. Remarks.-Shell form and penial shape Description.-Shell (Fig. 59) 1. 7-2.3 mm PROCEEDINGS OF T HE BIO LOGICAL SOCIETY O F WASHINGTON

,, ,

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10 ° 10 " 111"

Fig. 36. Map showing dimibulion of P. perturbalQ.

high, height/width, 140-190%. Whorls, 4.0- Outer lip thin, ncar-straight. Um bilicus nar­ 4.75, rounded, somewhat shouldered. Su­ row, chink-like. Protoconch mon ied wi th tures impressed. Body whorl 62-74% of shell series of shon, predominantly spira l stria­ height. Protoconch tilted relative to telco­ tions (Fig. 5ge). Teleoconch with wea kl y de­ conch. Aperture ovate, apertural plane veloped spiral li nes, panicularl y on upper slightly tilted relative to coili ng axis. Inner whorls. lip thin, only very slightly reHected, adnate Head/foot and visceral coil usually un i­ to or slightly separate from body whorl. formly coated with dark brown epitheli al VOLUME 102, NUMBER 1 m

Fia. 37. SEM micrographs of P cf. 5/rornsiana. USNM 853519. Sand Canyon (a. 3.04 mm; bar _ 136I'm). pigment. Central portions of sides of headl Central tooth broadly trapezoidal; basal foot consistine ofeilher a thin band or hroad eusps small. basal process only moderately patch, often lighter or unpigmented. excavated. La teral tooth having unusual Radular formula (Fig. 60): 4(5}-1-4/ 1(2}- "claw-like" appearance, with cusps few and 1, 2(3}-l-2(3), 16, 20 (from paratypes). widely separated. Peni s (Fig. 5 1b) elongate, '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY Of WAS HI NGTON

Fig. 38. Radula of 1'. cr. steGrapevine Mountains, Death Valley, Inyo Co., California. Distribulion and habitat. - Found in two thermal, highly mineralized springs o n limc:::otum:; bem;:h in northeast portion of Grapevine Spring complex (32-36"C, 1100- o 10SO micromhoslcm) (Fig. 52). Etym%gy.-Named after Peter Row­ lands, Resources Management Division, National Park Service, Death Valley, for his efforts in both assisting this study and con­ serving local aquatic gastropods in the Mon­ ument. , ,

Tryonia sa/ina. new species Cottonball Marsh tryonia Figs. Sic, 52, 61. 62 Tryonia sp.-Taylor in LaBounty and Dea­ o eon 1972:775.

Material examined.-Cali fo rnia: In yo Fig. 39. Reproductive anatomy of P cf. stearn· Co.; Spring in Cottonball Marsh (23 1), s;ana. USNM 857974, Sand Canyon: a, Len lateral USNM 860410 (holotype), -SS3556 (para. view of pallial ovid.ct complex; k. Peocs (b. c, dorsal types), 22 Feb 19S5; · 857998, II Mar 1987. aspects: d. e. ventral aspects). Diagnosis. - A moderate-sized species with elongate-conic shell bearing well-de­ veloped spiral striae. Penis with three pa· dorsaltcntacles, and all but narrow cen tral pillae on inner curvature and single papilla strip of head/foot. on outer curvature. Radular (Fig. 62) formula: 5(6}-1-5(6)/ Description. -Shell (Fjg. 6 1) 2.4-3.1 mm 2(3)-2(3), 3(4}-1-4(5), 19-26,26-28 (from high, height/width, 150-190%. Whorls, 4.5- paratypes). Central tooth broadly trapezoi­ 5.25, well-rounded, shouldered above. Su­ dal, basal cusps small , basal process weakly tures impressed. Shell loosely coiled, with cxcavated. Penis (Fig. Sic) longer than wide, slight separation of upper whorls common. moderately thickened. Distal ti p with Body whorl 6 1-68% of shell height. Pro­ prominent swelling of inner side. Inner cur­ toconch tilted relative to teleoconch. Ap­ vature wi th two small papillae positioned enure ovate, apenural plane slightly tilted on inner curvatllrc near tip, and larger basal relative to coiling axis. Inner li p thin, only papilla. Outer curvature with enlarged basal slightly reflected, narrowly adnate to or papilla. Dorsal penis variably pigmented by slightly separated from body whorl. Outer brown epithelial melanin, with especially li p thin, near-straight. Umbilicus chink·like dark cover on basal penis and papillae. to moderately open. Protoconch smooth Type localily.-Spring brook in Cotton­ (Fig. 6Ic). TeJeoconch with strong, regular­ ball Marsh, ncar base of Panamint Moun­ ly-spaced spiral striations, usually continu­ tainsat west si de of salt pan in Death Valley, ing onto body whorl, interacting with oc­ Inyo Co., Californ ia. casionally highly pronounced growth lines Distriblllion and habitat. - Found in a few to produce a weakly cancellate appearance. small , cool, bracki sh (17OC, >5000 mi­ Brown epithelial pigment dark on snout, cromhos/cm) spri ng brooks emerging in '" PROCEEDINGS OF THE BIOLCXilCAL SOCIETY OF WASHINGTON

. , ,. ., '. V

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,.. " " ". '.' • • • 00103050 ______~' Kllomo' . f .

117'

Fig. 40. Map showing distribution of P. cf. Sleornsian(1 in the study area.

Cotton ball Marsh, ca. 1.6 km out onto salt oped shell sculpture, penial loOO lian pat­ pan east of Salt Springs. Snails common on tern, and morphology oflateral radular leclh. large, actively growing tufa blocks. Highly remote area in ncar-pristine condition. Tryonia ~'ariegata Etymology. - Referring to highly miner­ Hershler and Sada, 1987 alized spring habitat. Amargosa tryonia Figs. 5 1c, 52, 63, 64 Remarks. - Vcry similar to T. rowlandsi from northern Death Valley, but dislin­ Tryonia variega/a Hershlcr & $ada 1987: guj~hcu \.Jy larger ~i./:c , ",uro,; Ioigilly lkvt;l- 817, VOLUME 102. NUMBER 1

Fig. 41. Camera lucida draw;n~ ofsllclls of P.... ong;: a. USNM 853531. sprinl alongside upper Pine Creck (2.18 mm): b. USNM 853524. spring N of Sum mil Cm:t (2 .24 mm): c. USNM, 853527. Hogback Creek (Lone PiM) (1.74 mm): d. USNM 853530. spring S of Warren Lake (2.08 mm)". e. USNM 853521. spring al Little Lake (1.44 mm): f. USNM 851947. French Spring (2.32 mm); g. USNM 851949, spring al ToU House (1.66 mm); II . USNM 857946. spring (n<)"h) al N end of Blind Spring Valle ~ (2.34 mm); i. USNM 853536. AnleloP<' Sprilll (1.'K1 mm): j. USNM 853535. Conal Springs (1.80 mm); k. USNM 853538. River Spri"&S (2.66 mm); I. USNM 853537. Layton Spring (1.78 mm).

Material examined. -California: San 1985; 857994, 22 Mar 1987.-Shoshone Bernardino Co.; Saratoga Spring (204), Spring (2 11 ), *853550, 12 Ma r 1985: USNM *853554, 27 Feb 1985; 857995, 10 857993, 16 Mar 1987.- Rcsling Spring Jul 1986. Inyo Co.; Spring crossing path on (212),853552,13 Mar 1985. south side of Amargosa Gorge (208), 853553 Diagnosis. -A variably-sized species with (empty shell).-Spring in marsh east of turriforrn-aciculate shell . Penis large, with Grimshaw lake (209), *85355 1, 13 Mar 3 or 4 papillae on inner curvature and single PROCEEDINGS OF THE BIOLOGICAL SOOETY OF WASHINGTON

Fig. 42. SEM micrograph. of P. wong; from weSI Sprilli in Birchim Canyon: a. Hololype. USNM 860403 (2.1 4 mm); b. e, Paralypcs, USN M 857941 (bar - 136 ~m). papilla (somelimes absenl) on oU ler cur­ 4.75-9.75, slightly to moderately rounded, vature. someti mes slightl y shouldered. Sutures im­ Descriplion.-Shcll (Fig. 63) 1.2-7.5 mm pressed. Body whorl 37--63% or shell height. high, height/ width, 170-350%. Whorls, ApenuTe nalTowly ovate, apertural plane VOLUME 102, NUMBER 1

Fi&. 43. Central radular teeth of p, wong!: a. USNM 857941. west sprint! in Birchim Canyon (bar - 10 "m): b. USNM 853521. spring at Liule Lake (bar - 7.5 "m): e. USNM 853536. Antelope Spring (bar - 7.5 "m): d. USNM 857981. River Springs (bar a. 12 "m). near-parallel with coili n& axis. Inner lip Umbilicus chink-like to moderately open. slightly thickened and reflected, adnate to Protoconch smooth to irregularly wrinkled or slightly se parated from body whorl. Out­ (Fig. 63d). G rowth lines often highly pro­ er lip thin, slightl y to moderately sinuate. nounced. 220 PROCEEDI NGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 44. L,lleral and rnarginallecllt or 1'. wQngi: a (laterals). e (laterals and inner marginals), d (ouler .nrginals), USNM 857~41. wesl spring in Birchim Canyon (bars - 6.0 I'm)" b. Laterals, USNM 853536, Antelope Sprin, (bar - 6.0 ~m). VOLUME 102. NUMBER 1 '"

j k

F" 45. Reproductive anatom~ of P. "wIg;: a, un lateral view of pallial oviduct comple~. USNM 857941. west sprinl in Birehim Canyon; b-I. Dorsal penes (b. USNM 857941 , west sprin& in Birehim Canyon; e. USNM 853530, spring SofWam:n Lake: d. USNM 857981 , River SprinlS; e, USNM 857979, Corral Springs: f. USNM 857980, Layton Spring: &. USNM 857947. French Spring: h. USNM 857949. spring al Toll House; i. USNM 8Sl534. spring (nonh) at nonhero end of Blind Sprin& Valley; j. USNM 857978. sprin& in Owens Gorge; k. USNM 857942. sprin& alonlSidt Lubkin Creek; I. USNM 853523. Summit Creek). ""c""-""I" drawn to same ~ ..

Radular (Fig. 64) formula: (4-7)-J-{4-7)/ black on snout and sides of head/foot, wi th 2(3}-2(3), 3(4)-1-4(5), 17-30, 22-33 (from lighter cover on central portions of latter. numerous populations). Central tooth typ­ Type/ocalily.-Five Springs, Ash Mead­ icall y broadly trapezoidal; basal cusps mod­ ows, NyC: County, Nevada. erate 10 large--si2ed, basal process moder­ Distribution and habitat. -Amargosa ately eltcavated. River drai nage, from Ash Meadows south Epithelial pigment usually dark brown- to Saratoga Spring. Snails typicall y common PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHlNGTON

f

,

;

I"i&. 46. !'ene. (ventral a.pcct.) or P. wo"g;: D, USNM 857941. west $pr;ns in Birchim Canyon; b, USNM 857947. Fmtch Spring; c. USNM 853536, AnteLope Sprin&; d, USNM 857942, spring alongside Lublin Creek; e. USNM 853534, spring (no"h) at no"hem end of Blind Spring Valley; f. USNM 857981. River Sprinp; L USNM 857949, spring at Toll House: h. USNM 857979, COmll Spring;;. USNM 853523. Summil ~k. 00.-­ 'Oe" and "8"_0.;" drawn 10 same scale. on various substrates in small springpools wongl): and c) small-sized Tryonia spp., in­ and upper portions of oudlows (7-32OC, cluding those of northern Death Valley (T. 430-10,150 micromhos/cm). salina, T. rowlandsi, T. margae) and Ash Remarks. - Populations from lower Meadows (T. ericae, T. elala). Amargosa River drainage are clearl y as­ Results (summarized in Tables 3-5) sup­ signable to T. I"ariegala on basis of shell , port taxonomy presented herein , insofar as radular, and penial features. high correct classifications (81-87% lover­ all» indicate that these snails are well-dif· feren tiated in shell fealures. In the analysis Morphometries involvi ngP. amargosaeand P. micrococcus, Discriminant analyses were performed on correct classification was 89% (40/45) and each of three groups of congeners from giv­ 80% (29 1/362) for each species, respective­ en portions of the study area: a) P. amar· ly. Correct classification for two populations gosaeand P. micrococcus. very similar forms of P. amargosae (Amargosa Gorge, Grim­ with cOnliguous ranges; b) Pyrgulopsis fa u­ shaw Lake) from Tecopa area (where the na of O",ens Valley (P. aardahli, p, owen­ ranges abut) was high (83%), as was that sensis, P. perturbata, P. cf. stearnsiana, P. (92%) for two nearby populations of P. mi- VOLUME 102, NUMBER I m

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'0 0 ' 0 30 ~O ___ ~ __~ , KilO"".'. , 11'· , 17· Fig. 41. Map showina dislribulion of P.... ongl. crococcus (Tecopa Hot Springs, Shoshone In this analysis, discrimination was based Spring). Misclassified specimens of P. mi­ largely on height of shell and aperture. crococcus were concentrated in a few pop­ For the Owens Valley Pyrgillopsis. correct ulations in nonhern Death, Panamint, and classification .....as 84% overall (382145 1), Saline Valleys (well separated from the range and ranged from 79_87% by species. The of P. amargosae), and indicate local differ­ sole species having <80% correct classifi­ entiation of this relatively widespread form. cation, P. owellSensis. was most frequently PROCEE DINGS OF THE BIOlOGICAL SOCIETY OF WASHINGTON

Fig. 48. SEM micrographs of T. margaI" a-

Fig. 49, Central and latcral radular teeth of 7: mQ,gQ~: a. c. USNM 857953, Grapevinc Springs. upper warm spring (a. centrals. bar - 6.7 "m: C, latcrals and inner marginals. bar - 6,7 "m): h. d. USNM 857996, Grapevine Springs. spring abo~e shack (b. centrals. har - 6.7 "m; d. laterals, har - 6.7 "m).

(10/ 14) misclassiftcd as P. wonKi. which nant runction were related to size and width cannot be conrused with the rormer when or shell and aperture. penial morphology is considered. Parame­ Classification or each or the three small­ ters weighing heavily on the first discrimi- sized Tryonia spp. from northem Death PROCEEDINGS Of THE BIOLOGICAL SOCIETY Of WASHINGTON

fig. 50. Margi nal radular (CClh. of T. ",,,rg"t>: a. c. USNM 857953, Grapevine Springs. upper warm spring (a, lau,rals and inner marginals. bar - 10.0 I'm; C, OUler marginal. bar - 3.8 I'm); b. d, USNM 857996.Grapevine Springs. spring above shack (b. lalerals and inner marginals. bar - 7.5 I'm: d. outer marginal. bar ~ 3.8 I'm).

Valley was 100%, while that of T. elata and were well-discriminated, with only a single T. ericae from Ash Meadows was 93% and specimen (of the lattcr) misclassificd (as the 95%, respectively, Despite their general former). Parameters weighing heavily on thc similarity in form, T. margae and T. elata fi rst discrimi nant function were shell height, VOLUME 102, NUMBER t m

Table 3. - Resulls of discriminam fUrK:I ion analysis on Pyrgulopsis arMrgoslU' and P. m;cl"O<'O<:t'W. Also P"cn are the eancnical oolRlat;on. number of speci­ menS used (by species. ordered as above). and perttll1 COrT«t classi fi cation (overall).

SWIdafd,.ed V, ,,,,bIn ,,,,,II><> C""',.. ,"", WH 0.365 0.554 SH 1.1 94 0.354 SW 0.398 ~0. 09 4 E L.W - 0.717 0.176 E w.w 0.749 0.] 46 Q AL -I. 728 0.055 AW - 0.067 -0.108 W 0.176 -0.292 D 0.798 0.536 T -0.390 0. 425 SA 0.87] 0.351 C. oorrela tion 0.475 N 45.362 'iii corre<:! d anificatioll " F".. SI. Penes (dorsal aspectS) of TryOllia spp.: a. T. 1fIlUgtIe. USNM 8S19S2. Grapevine Springs. upper wann ,prina: b. T. ro ...la lldsi. USNM 857953. Grape­ 1978; Smith & Street· Perroll 1983; Minck· vi!leSprinp. vpperwarm sprina: c. T. '"Qn~a. USN M ley et al. 1986; Taylor 1986). Configuration 8S7 994. spri"l at Grimshaw I.ake: d. T. rOOus/a. USNM of the System i>; ponrayed in Fig. 6S as pre· 8S7999. Nev~ Spri ngs. upper spri ng; e. T. salioo. USNM 8S7998. Cononball Marsh. "a." "b." "d," "e" lude 10 discussion of springsnail zoogeog· drawn to same scale. raphy. Springsnail d istributions re fl ect pro· nounced local endemism of the modem rau· width of body whorl , and apenure size and na. Twenty of 22 species known from Death shape. Valley System u ea are restricted 10 its con· fin es. Of these, 10 are endemic to single springs or spring complexes, and another Discussion scven occur in relatively small portions of Modem drainage in the Death Valley area single modem drainage basins. Locally en­ includes Owens, Amargosa, and Mojave demic Pyrgufopsis are concenlrated in Ow. Riven , the laner two of which are dry along ens Valley and Ash Meadows, whileendem­ most of their courses. Allhough now iso· ic Tr)/onia occur in the laller and non hem lated from one another, these river systems Death Valley. Three forms are relatively were integrated during wener, pluvial times widespread in the system: P. wongi.of plu. during the Pleistocene as a seri es of lakes vial Owens River drainage (Long, Adobe. overflowed along their courses 10 terminate Owens Valleys); P. micrococcus. wi dely dis· in Death Valley. This Death Valley System tributed from Amargosa Valley west 10 Pan­ (named by Miller 1943:69) was initiall y amint Valley; and T I'ariegata, of pluvial proposed by Gale (1 9 14) and Blackwelder Amaraosa Rive r drainage (Ash Meadows. ( 1933), and has been discussed in detail by Tecopa Basin, Sar3l0ga Spring (southern many others (Hubbs & Miller 1948; Miller Death Valley]). 1946, 1948; Smith 1978; 50hz & Naiman Endemic p)'rg!llopsis ofO... .. ens Valley and 228 PROCEEDINGS O F T HE BIOLOGICAL SOCIETY OF WASHINGTON

, , " T. margae c T. prot ee 4 T. ro buSl a •

~ . T. r owland s i 0 T. salina t. , T. variegala . " ,~ ' ' j "I.( - rf'!- -. , ••

•

Ash Meadows

• ., 0 " • "" ". " " -. '-' •• • NI " ,-. ,", . , • • b '-J .. .. • " .., K llo",e'." " a' " ,. Fig. S2. Map showing distribution of Tryonjo. spp. in study area. Tyronia .wif'gtlla occurs al I i localities in Ash Meado ....'s.

Amargosa Basin arc sharply differentiated Amargosa drainage), include a group com­ on basis ofpcnial morphology and represent prising P. cr. slearnsiana and similar, en­ separate local radiations. Owens Valley demic forms (P. aardahfi. P. oll"ensensis, P. forms, characterized in part by possession perturbata); and P. wongi, which has no close of a ventral penial swelling bearing glan­ relatives in the study area. Ash Meadows dular ridge (abscnl in forms endemic 10 (Amargosa Basin) endemics represent at VOLUME t02. NUMBER I

Fia,. SJ. SEM micrographs of T. pro/ea. USNM 857954. HOI Creek. StleU ··a" is 3.64 mm lall. ··b." ··c·· show sculplure on prolOCOnch of embryonic shell (bars - 60. 17.6 "m). least two lineages (both including Fllllllin­ banksensis. P. nallllS. P. iso/milS). The latter ieo/a-like representatives): snails with re­ may be related to the probable sister species duced or absent penial lobes and a glandular pair of widespread P. micrococclis and P. ridge on mid-ventral penial surface (P. amargosae from southern Amargosa River er},thropoma, P. crysra/is. P. pisrerO; and drainage. those having a ridge on terminal ponion of Relationships among regional Tryonia are moderate-large sized penial lobes (P. fair- more speculative. Endemic Tryonia in

Table 4. - Results of discrimina~1 function analysis on O,,-cns Vallcy Pyrtu10psis (P. aa,daMi. P. o"en5en5is. P penU,/JQIQ. P. cr. 5IeQmsia·e). and percent correcl classification (o>waU).

Sut>n

Fig. 54. Radula of T. pro/eo., USNM 857954, HOI C,,,,,k: a, Cemrals (bar - 12 I'm); b, c, Lateralsand inner marginal. (l>ars - 8.6 I'm); d, OUleT marginal. (bar - 7.5 "m). VOLUME 102, NUMBER I 231

F;" ~~ . SEM micrographs of T. robus/a from Nevares Spri ngs. upper spring: a. Holotype. USNM &604 11 (1.92 mm); !>-d. Paratypes. USNM 853557. nonhern Death Valley and Ash Meadows, tative from the east, T. \'oriegol o (contrary apart from enigmatic T. robusUJ, apparently to Taylor 1985: 3 17) and may not represent comprise a single local radiation. T. proteo a separate lineage (as implied by Taylor from west of Death Valley (Long and [fossi l; 1985:3 17). Taylor 1985:317) Panamint Valleys) is quite Pluvial waters of the Death Val ley System similar (in shell ) to at leasl one represen- provided aquatic continuity (and thus the

Table 5.-Results of discriminant function analysis on small -sized Tryon!a (T. ela/a. T. ericae. T. margfle. T. ro ... /aoosi. T. salina). Also give ~ arc canonical correlations, number of specimens used (by spe<:ies. ordered as above). and per<;ent correct classification (overall ).

SUn4a

Fig. 56. SEt.! micrographs of T. ,ohm/a. Nevares Springs, upper spring, USNM 853557: a-<:. Views ofaP9 (ba["l; - 75. 43. ]00 I'm): d. Dorsal o perculum (bar - 200 I'm). VOLUME 102. NUMBER 1 233

Fig. 57. Radula of T. robU!>la. Ncvares Springs. upper spring. USNM 857999: a. Ccmrals (bar - 6.0 JIm): b. C. Lalerals and inner marginals (bars - 3.8, 5.0 JIm): d. OUler marginals (bar - 3.8 JIm). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 58. SEM micrographs of penes of r . robu$la. USNM 857999, Nevares Springs. upper spring: I. Dorw­ righl laleral aSpeCl. wilh allachment 10 head on lower left (bar - 136 jim); b. Distal lip of papilla (N., - 23.1 jim); e. d. Dislallips of penis. showing ciliation (bars ~ 30. 20 jim). VOLUME 102. NUMBER I

e

Fi&. 59. SEM micrographs of T. fOK·!tmdsi from Grapevine Sprinas. upper warm spring: a, HolO1Ype. USNM 860409 (2.18 mm): b-e. Paral},l)eS. USNM 857953 (bar - 100 I'm). 216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHI NGTON

fig. 60. Radula of T. ro .. la"dsi. USNM 857953, Grapeville Springs, upper warm spring: a, Celllnis (bar ­ I S I'm): b, Lalcral. (bar ~ 13.6 jJm): c. Inner marginal (bar - 6.0 I'm): d, OUler marginals (bar - S.O ,..m), means for potential exchange ofaquatic fau­ em springsnail zoogeography indicates tha i na) across current drainage divides imposed the above did not facilitate much mixing by the region's predominant north-soulh between pluvial Owens and Amargosa Riv­ trending basin and range topography. Mod- er faunas. These results support earlier con- VOLUME 102, NUMBER I 237

fl&. 6 1. SEM , " i ~ l U!!1a vh. uf T. ,,

a1l Manh. II. H olo!)!>". USNM 860410 (3.27 rnrn); b, c. Paratypes. USNM 853556 (bar - 120 I'm).

dusionsofTaylor (i n Miller 198 1: 58, 1985: vial la kes have not becn thoroughly inves­ 317- 3 18), although his observation that tigated (see Hershler and Minckley 1986). "Spillover from Lake Panamint into Death Details such as size, number, and salinity Valley ... had no recogn izable effect on ofinter-connected lakes; and length and gra­ mollusc distribution" (Taylor 1985:3 18) is dient of lake outlets obviously would affect contradicted by distribution of P. micro­ probability of snail dispersal. Further, other coccus (and possible close a ffi nities between modes of dispersal undoubtedly occurred T. proteo and congeners to the east). and could have produced distributions Taylor ( 1985:317) suggested that faunal comparable to or conflict ing wi th those pre­ exchange between Amargosa and Owens dicted by pl uvial drainage models. The River drainages might have been precl uded presence orraun a in isolated basi ns rt.""(:ently by a saline dispersal barrier consisting of an created or devastated by volcanic activity arm of Bouse embayment (from fonner G ulf (Long, Adobe Valleys, respe<:tively), or those of California) extending into Death Valley. that either lacked pluvial connections (Sa­ Although this hypothesis appears plausible, line Valley, Frenchman A at(Cane Spri ng]; it may not need to be invoked to explain Hubbs and Miller 1948) or connected with patterns described above. Apart from ad­ snailless areas (Deep Springs Valley, whose d itional inHuences on snail distribution im­ pl uvial la ke spilled into Eureka Vall ey; Mil­ posed by fa r older lacustrine episodes than ler 1928) attests to importance of factors considered above (Smith 1984) and other other than late Pleistocene pluvial drainage aspe<:ts of complex regional geologic/hydro­ systems in effecting modem snail distribu­ logic history (Minckley et al. 1986), fu n­ tions in the region. A detailed zoogeograph­ damental questions rela ti ng to types ofhab­ ic analysis encompassing the above consid­ ita! occupied by Pleistocene springsnails and erations will be provided following plausi bility of dispersal of such (frequently completion of springs nail survey of remain­ stenotopic) organisms along chains of plu- ing portions of the Death Valley System. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 62. Rad",la of r. salina. USNM 857998. COllonball Marsh.: a. Centrals (bar - ]0 I'm); b. laterals and inner marginal (bar _ 7.5 ~m): c. Inner and Outer marginals (bar - 8.6 I'm); d. Outer marginal (bar - 5.0 I'm). VOLUME 102, NUMBER I 2"

Fill. 63. SE M micrograph $ of T. mriegal(J: a (2.S mm). b. USNM SH554, Saratoga Spring: c (2.S6 mm). d. USN M S53550. Shoshone Spring (bar - 86 jim). Fig. 64. Radula of T. l'lJriegata: a. b, USNM 857995, Saratoga Spring (a. ""ntrals, bar - 10.0 I'm: b, laterals. bar - 7.5 I'm): c, d. USNM 857993. Shoshone Spring (c. laterals and inner marginals. bar - 6.0 I'm; d, outer marginal. oor - 4.3 I'm).

Fig. 65. Map of Pleistocene Dealh Vallcy System showing drainage relations (not necessarily contempOra­ neous), Stippled areas encirCled by dashed lines may nOI have oontained lakes for significant pOrtions of tbe pluvial period. In Ihe northeast, Lake Russell (Mono Valley) spilled into Lake Adobe (Adobe Valley). which in !Urn o"crAo",cd (Ihrough a gap in Benlon Range) 10 enter Ihe (now mostly dry) north fork of ancestral Owens River (Putnam 1949: Hubbs and Miller 1948: Gilbert cl al. 1968). La ke Long Valley filled shortly afler Long Vallcy caldera " 'as created by cxplosivc eruPtion ofUishop luIf700.000 BP (Bailey et al . 1976). and spilled inlO south fork of Owens River. OUIAo\\' from Owens Lakc, pOssibly initiatcd as a result of increased How due: \0 '" ,

,.... ,..

o 25 50 75 LI __JI L _ --,I __-,I Kilometers

".

,. 0 ••••

,. ....' ,

,. ••

B ase map h om S nyder et 31 .1964 1 1 8° 'I So capture or upper San Joaquin River drainage in u mg Valley (Huber 1981 , Smillt el 31. 1983). passed beN-cen Sierra N",-ada and Coso Range 10 erner and fill China and Searles Lakes. Searles Lake overftowed ;1110 Panamim Valiey. which held a large lake in its southern ponion that spilled into Death Valley during several periods be,,,-ecn 120,000 and 20.000 BP (Gale 1914. Hooke 1972. Smith 1976. Hale 1984). To Ihe ,,-eS!. the lush wetland of Ash Meadows (lha! did nOI have a large pluvial lake: Mifflin and Wheal 1979. Hay et a1. 1986) drained 31 leaSI imcrmillemly 10 Amargosa River during Pliocene· Pleistocer.e. Funher downAo,,' Ihe ri "cr was ponded In mid- 10 Jatc-PleislOcc n{' by alluv;al fans ab(l,·c Amafll,osa Gorge 10 form Lake Teopa (Sheppard and Gude 1968. Starkey and Blackmon 1979). which spilled imo southern Death Valley. Moja"e River tined a series of three pluvial la kes. wilh Lake Moja,·" o"crBowinlltOward soulhern Deat ~ Valley during several intervals of lale Pleistocene from > 14.500-9000 BP (Ore and Warren 1971). Overflow from Dealh VaUc}' S}'slem 10Coiorado River duri ng an e ~ lreme pluvial period is conje<: lural. and has often been hypothesized based on dislribulion ofaqU3tie biOla (Miller 1981. Taylor 1986). Ha le (1984) dcscribed an old (pre-Pleistocene) fluvial channel al Ash Hill near l udlow. CA. lhal could be a produci of such oUIHow. The channel is al 594 m elevalion. which would require a "aSI pluviallakc ( > 300 km long and > 650 m maximum depth) extending 10 northern Death Valley, additional geological "vidence is needed to conform this rem arkable possibilily. '" '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments M"'alt;a. _ Procttdil\&S of the Academy ofNal_ ural Sciences of Philadelphia 7:269. Field"'ork in the Dealh Valley area was Gale. H. S. 19 14. Salines in lhe Owen$, Searles. and supported by Smithsonian Institution Re­ Panamin Basins. southeas":rn California.­ search Opportunity Fund (ROF), Wildlife United States Geological Sur... ey Co:nrib'l1ions Conservation In ternalional orthe New York to Economic Geology (1913):25 1-3B. G ilben. C. M .. M. N. Christensen. Y. AI- Raw;. & K. Zoological Sociely, and contracts awarded R. Lajoie. 1968. Structural and ... okanic his­ by U.S. Fish and Wildlife Service, Great lOry of Mono Basin. Califomia-Nevada._Geo­ Basin Complex (Order No. 14320-0 182); logical Society of America Memoir 116:275- Bureau of Land Management, California 329. Desert District (CA-060-CTS); and Califor­ Gould. A. A. 1855. New species of land and fresh­ water shells from ,.·eslern America.-Procttd­ nia Fish and Game (C-1 922). Collecti ng ings of the Boston Society of Natural History 5: permits were provided by National Park 127-130. Service (Death Valley Nalional Monument) G regg. w.O.. & D. W. Taylor. 1965. ;olltrll«l/a and Siale of California, and access to sites (Prosobranchia: Hydrobiidae). a new 1CIlU$ of on Nevada Test Site, China Lakes Naval West American freshwalCrsnail •. _ MalacoJogia 3:103-110. Weapons Center, and Fort Irwin is also Hale. G. R. 1984. Reassessment of the Death Valley_ gralefully acknowledged. Bureau of Land Colorado River overflow h~'polhesis in light of Management (Distri ct o ffices in Ridgecrest new evidence. Ph.D. dissertation (unpublished). and Bishop), California Fish and Game Unive.. ity of California. Berkeley. 291 pp. (Bishop office), and Nati onal Park Service Hay. R. l.. R. E. Pexton. T. T. Teague, &, T. K. K~ r . 1986. Spring-relatcd carttonatC rocks. Mg days. (Death Valley National Monument) loaned and a~wc i ated minerals in Pliocene Great Basin. wi th emphasis parameters. on Glacial and Poslg)acialtimes. Bullelin of the Conrad. T. A. 1855. Description ofa new species of Universi1y of Utah 38. Biological Series 10. VO LUME 102, NUMBER I

Huber. N. K. 1981. Amounl an.d timing of laIC Ce­ and &Cnnis of aUlhiacnetic silicate minerals in nozoic uplift and tilt of the Central Sierra Ne­ ,uKs of Pleis:ocene Lake Tecopa, In)"o County. vada. California _ evidence from the Upper San California. _ United States Geological Sun'ey Joaquin River Basin. - Unittd States Geological Professional Paper 591: 1-38. S u ~ Professional Paper 11 97:1-28. Smith. G. I. 1984. Paleohydrologic regi mes in the Kohn. A, J .. &: A. C. Riggs. 1915 Morphometry of soulhweslem Great Basin, 0-3.2 my Ago. com­ the CO" IIJ $hell.-Systematic Zoology 24:)46- pared wilh other long records of "global" cli· )59. mate.-Quaternary Research 22: 1-17. LaBounlY. 1. F .. &: J. E. Deacon. 1972. Cyprmodon --. V. 1. Barczak. G. F. Moulton,&:J.C liddicoat. millen, a new species of p~ pfish (Family (:y­ 1983, Con: KM -3. a surface-to·bedrock record prinodomidae) from Death Valley, Califor_ of Latc Cenozoic sedimentation in Searles Val­ nia. _ Copeia 1972:169-780 ley. Californ:a.-United States Geological Sur­ Milftin . M. D .. &: M. M. Wheat. 1919, Pl uviallakes vey Professional Paper 1256:1-24. and estimated pluvial climales of Nevada. - Ne­ --, & F. A. Sm ci-Perroll. 1983. Pluvial lakes of vada Bureau of Mi ~ s and Oeology 94: I-57. the United S:.ales. pp. 190-212 in H. E. Wright, Miller, R, R. 1943. C}'Prinodon soJinu$, a new species ed .. Latc-qllalcrnaryenvironmenlSoflhe United offish from Death Valley, Califomia.-Copeia States, Vol ume I. The Latc Pleistoccne (S. C 1943:69-78. Porter, ed.). University of Minnesota Press, --. 1946. Correlation bet,""""n fisll disuibution Mi nneapolis and Pleistocene hydrograpt,y in eaSlern Cali­ Smilh. G. R. 1978. Biogeography of intcrmountain fornia and soutllweslern Ne"ada. wilh a map of fishes. pp. 17-42 in K. T. Harper and J. l. Re­ the Ple,stocene waters.-Joumal ofv eology ~4: veal , eds., Intermountain blogeOgraphy. a sym­ 43-53. posium. Grell Basin Naturalisl Memoir 2, --. 1948. The cyprinodonl b hes of the Death Smith, R. S. 1976. Lale_Quaternary pluvial and tec­ Valley System of eastern California and soUlh_ tonic history of Panamint Valley. In)"o and San western Nevada.-Miscellaneous Publications Bernardino Counlies, California. Ph.D. disser­ ofthe Museum of Zoology. Un iversilY ofMich_ tation (unp~ b ljs lled ) , Cal ifo rnia Inslitule of ;pn 68: I-ISS. Technology, 295 p. --. 198 1. CoeVOlution of desen and pupfishes Snyder. C. T, G. Hardman, &: F. F. Sdenek. 1964. (genus Cyprinodo,,) in the American Southwest. Pleistocene la kes in the Greal Basin.-United pp. 39-94 in R. J. Naim an and D. l. Soltz. eds.. States Geological Survey Miscellaneous Geo­ Fishes in Non h American dcsen s. 10hn Wiley logical InvcS/iga,ions Map 1_416. and Sons, New York. Soltz, D, L. , & R. J. Naim an. 1978. The natural --, &: E. P. Pister. 1971. Management of tile history ofnalive fishes in the Death Valley Sys­ Owens Pupfish. C),pri"odon , adiosus, in Mono tem ,-Nalural History Museum of l os An8eles CounlY, Cali fornia.- TranS3Clionsofthc Amer· County. Science Series 30:1-76. ican Fisheries Sociely 100:5[12-509. Starkey. H. C. &: p, D. Blackmon. 1979. Cia)' min_ Miller, W . J. 1928. Oeology of Deep Spring Valley. eralogyofPte!stocCne Lake Tecopa. Inyo Coun. California. _ Journal of Geology 36:51()"'S25. ty. Californi •. -United States Geological Sur. Mirw,;kley, w. l.. D. A. Hendrickson. and C. E. Bond. vey Professional Paper 1061: 1-34. 1986. Geography ofweslem Nonh Amencan Steams, R. E. C. 18 93. Repon on land and fn:shwater fn:shwater fishes: Descriplioo and relationships shells collected in California and Nevada by the to in lracontinentaltN:tonism. PI'. 519-613 in Death Valley E~ pedilion, including a few ad· C. H. Hocull and E. O. Waey, eds.. Zoogcog­ ditional spc<:ies obtained by Dr. C. Han Mer. I"\lphy ofNonh American fremwaleT fis hes. John riam and assistants in pans of the south,.'estern Wiley and Sons, New York. Uniled States.- Non h American Fauna 7:269- Ore, H. T., &: C. N. Warren, 1971, Lale Pleislocene­ 283. Early Holocene 8eomorphic hislory of Lake Mo­ Taylor, D. w . 1950. Three new l'yrgulopS/S from the jave, Cali fornia.- Geologictl Socie,y of Amer­ Colorado o.""n. California.-lcaHets in Mal. ica Bulletin 82:2553- 2562. acology 1:27 ~ n , Pilsbry. H. A. 1899. Catalogue of the Amn;colidae --. 1966. Sunmary ofNonh American Blancan of the Western Uni led States.-Nautilus 12: 121- nonmarine mollusks.-Ma!acologia 4:!_172. 127. --. 1981. Frnh,.-aler mollusks of California: A Putnam, W. C. 1'>4'>. Quatemarygcolo&y of'he June di.uib",ion.1 eheckli".-Califomi" Fish and Lake DiSirict. Califomia,- Bulletin oflllc Geo­ Game 67: 14)...163, logical Sociely of America 60:1281-1302. --. 1985. Evolution of freshwater draina8es and S~ pa rd , R. A.. &: A. J. Gude. 1968. Distribution molluscs in western Nonh America . PI'. 265- '" PROCEEDINGS OF THE BIOLOGICAL StX:IETY OF WASHINGTON

321 m C. 1. Hocull and A. B. Le"IlOll. t'ds .. La ~ CA. 12.6 km S-SW of NE com.... of quad.ran&le_ [586 CenOlok HiSlor), of the Pa d~c Nortbwest. m. 4·30-87. 19. Sprincs in (2) unnamn! ~o n s N of American ASSOC;'lion rOT the Ad"ancement of Haiwee Canyon. CA: Inyo: Monaclte Min.. CA. NW Science. San FranciK<), California. comer.; sees. 30_ 3 1. T 20S_ R 37E. 1556 m. 4-30-87. Wilkinson. l. 1986. SVST AT: The system fo r s .... • 20. Hogback Crerk. CA: Inyo: Monache Muo .. CA_ 7.3 tiSli~. SYSTAT.lnc.• EvansIOIl.IL. km SW ofNE oorn.... ofquadranpe. 1586 m. 21. Sum­ mit Crerk. CA: [nyo: Monachc Min .. CA. SE V. sec. Depanment of Invertebrate Zoology, 7. T 2OS. R H E. 1313 m. 22. Spring ca. 3.0 km N of Summit C=k. CA: Inyo: Monachc Min. CA. CCIItn" NHB STOP 11 8, National Museum ofNal­ of sec. 6. T 2OS. R 31E. 128] m. 23. WaI~ Cfttt­ ural Hi story, Smithsonian Institution, CA: In)"o; Monache Min .. CA. NE 110 sec. 34. T [95. Washington. D.C. 20560. R 37E. 1769 m. 4-16-87. 24. Carugo C.....,k_CA: Inyo: Olancha. CA. NE V. sec. II. T [ 9S. R 37E. 1159 m. 4-16-87. 25. Sprinlat Cabin Bar Ranch. ca. 2.S bn N Appendix of Olancha. CA: Inyo: Olancba. CA. SW \\ sec. 6. T Collection localities. numbemi as in Fits. 2-7. Da .... 19S. R 37E. 1098 m. 26. BraleyCreek and s~npjl.lSl include name orsil". stale. counry, topographic sheet. lOS. CA: Inr o: Olancha. CA. 10.Okm NW ofSEo;omer IOWTlship aid rang!! coordinates, site elc"I lion. and of quadrangle. 1100 m. 4-16_87. 27. Sprinp on cd&t: dale of "isitalion (for negative si tes o nly). ofo-.."Cns Lake at Permancntc. CA: Inyo: Olancha. CA. 13.8 km N of SE comer of quad.ran&le. 1068 m. 2-9- I. Su-eam in Saat: Canyon. CA: Kern; Horse Canyon. 85. 28. Ash C=k. CA: Inyo; Olancha. CA. 15.6 km CA (7.5), 7.0 km sw orNE comerofquadran&le, 1342 NW ofSE comer ofquadran&l(. 1068 m_ 4-16~ 7. 29. m. 2. Boulder Spring. CA: Kern: Horse Canyon, CA. COllon""QOd C=k. CA: Inyo: Olancha. CA. 9.6 bn S.6 km S-SW of NE romer of quadrangle. 1251 m, sw ofNE comer of quadrangle. 1037 m. 4-16-87. 10. 3-26-87.3. Stream in Indian Wells Canyon. CA: Kern: Lo"'.... CenlOmnial Spring. CA: Inyo; Keeler. CA. 2.3 Inyokern. CA. NW If. sec. 17. T 26S. R 38E. [068 m. km W of SE comer of quadrangle. 1769 m. 3-31-87. 3-26-87.4. Sprillj in SW COmer ofSllon Can}·on. CA: J I. Oirw Socks (HOI Sprinl). CA: In)"o: Keeler. CA. Kern: [n)"okern. CA. NW If. sec. 5. T 26S. R 38E. 1129 NE 1'. sec. 34. T 18S. R 37E_ 1098 m. 2-9-85. 32. m. 3-26-87. 5. Stream in Grapevine Canyon. CA: Kern: Springs at S end of o-..=s Lake. ca. 3.5 krn NW of Inyokern. CA. cenler of .....,. 29. T 25S. R 38E. 946!1l. Diny Socks. CA: Inyo: Keeler. CA. NW .... occ. 11. T 3-26-87.6. Stream in Sand Canyon. CA: Kern: Linle 18S. R 38E. 1098 m. 2-9-85. 33. Sprincs on cd&t: of Lake. CA. center of sec. 1. T 25S. R 38E. 1068 m. 1. Owens Lake. ca. 3.0 km S of Keeler. CA: In\'o; Keeler. Stream in Noname Canyon. CA: Kern: Little Lake. CA. NW 110 sec. 22. T 17S. R 38E. 1098 m. 4-25-87. CA. 10.0 kill NE ofSW comer of quadrangle. 916 m. 34. lubkin Creek and spring fcedingcreek from _ lb. 3_26_87. 8. Stream in Ninemile Canyon. CA: Inyo: CA: In )"o: Lone Pine. CA. SE 1'. sec. ]6. T l.s. R 3610. little Lake. CA. ]2.2 km NE of SW comer of quad_ 1220 m. 35. Spring along E side ofTutlle Creek. CA: rangle. 916 m. 3-26-87. 9. Sprillj 0.8 km S of Little In)"o: Lone Pine. CA. NE If. sec. 6. T 16S. R 36 E. 128 1 lake. W of HW 395. CA: In)"o: Little Lake. CA. SE If. m_ 36. Hogback Creek. CA: In)"o: lone Pine. CA. NW sec. 18. T as. R 38E. 946 m. 4- 1-81.10. Spring at 't. sec. 2. T ISS. R 35E. 1159 m. 37. Sprina II NE end litt]c Lake. E of HW 395. CA: In)"o: little Lake. CA. of Alabama Hills. ca. 4. 2 km N_NW ofl..one Pine. CA: NW \4 sec. 11. T 23S. R 38E. 946 m. II. Siream in In)"o: Lone Pine. CA. NE '/. sec. 31. T 14S. R 36E. Linle Lake Can)"on. CA: In)"o: Linle Lake. CA. NE If. 1159 m. 4-25-87. 38. George Creek_ CA: Inyo; loDe sec. 12. T 2JS. R 37E. 1129 m. 4- 1-87. 12. Springs ca. Pine. CA. NE \\ sec. 27. T 14S, R 35E. 1251 m. 4-25- 1.0 km N of LinIe LakeCanyon. CA: Inyo: Little Lake. 87.39. Independence Creek. CA: In)"o: I n~_ CA. SW 1'. sec. I. T 23S_ R 37E. 1159 m. 4-30-87. 13. CA. SE If. sec. 23. T US. R 34E. 1342 m. 4_17_8 7. 40. Slream in canyon ca. 3.0 km N of lillIe Lake Canyon. Boron Springs. CA: In}'o: MI. Pincllot. CA, NW .... sec. CA: In)"o; Linle Lake. CA. 4.4 km SE of NW comer 22. T 13S. R 34E, 1556 m. 41 . Oak Creek. _th fort. ofquadrantle. 1220 m. 4-30-87. 14. Stream in PoT- CA: [nyo: MI. Pinchot. CA. SW 1'. sec. 10. T IJS. R 1ug<:Se Canyon. CA: 1nyo; Hai wt'C R~r,"oir. CA. 2.19 34E. 1525 m. 4_1 7_87. 42. Sprinp ca. 1.0 bn W of km NE ofSW comer of quadrangle. 1342 m_ 4_1_81. M1. Whilney Fish Hatchery. CA: In)"O: ML PinchO!. ] 5. Sprinp on Pan"",.., Iknell. CA: Inro: Haiwee CA. SE If. sec. 3. T 13S. R 34E. 1342 m. 4_18_81. 43. Rese .....·oir. SW COmer.; ofsecs. 3. 10. T 22S. R 37E. Siream in Charlie Canyon. CA: In}'o; Mt. Pir.cho!. CA. ] 160-1220 m. 4-30-87. 16. Lo""Cr spring in Tunawec SW II. sec. 3_ T IJS. R 34E. 1617 m. 44. Springs fecd..in& Canyon. CA: [nyo: Haiwcc R~ ..... oir. CA. SW \4 sec. N fork Oak Creek. CA: In)'O: Mt. Pinchot. CA. SW V. 33. T 2]S. R 31E. 1313 m. 4_25_87. 17. Upper spring sec. 3, T 13S. R 34E. 1586 m. 4-25-87. 45. Oak Creel<. in Tuna"-,,e Canyon. CA: Inyo; Monache Min., CA. nOllh fork. CA: Inyo; MI. Pinchot. CA. <:enler of sec. 1.0 bn N_NW of SE com er of quadJan&le. ] 525 m. 3. T 13S. R 34E. ] 525 m. 4_1 7_87. 46. Grov.... Antoo 4_30-S7. tS. Haiw-ee C.....,k. CA; Inyo; Monache MIn .• Spring. CA, Inyo; MI. Pi n~ho t . CA. SW V. "'" 20. T VOLUME 102. NUMBER I '" us. R :WE. 1586 m. 4-18.87. 47. Harry Birdl Sprinas. SIo"",. "NE Spri~g." CA: Mono: While MIn. Peak. CA: Inyo: MI. PinehOi. CA. NW >;. i«. 17. T 12S. R CA. NW h sec. 18. T 5S. R BE. 1281 m. 78. Anlelope :WE. 1464 m. 4.18.87. 48. Sawm~1 CIftk. CA: In)'o: Spring. CA: Mono: Casa lM.blo Min .. CA. SW ~. sec. MI. Pinehol. CA. SW \'. sec. 9. T 12S. R :WE. 1251 9. T 4S. R l iE. IHO m. 5-1·87. 7'1, M o~n Spri"l. m. 4- 17.87. 49. Oiv;S!on Creek and Scott)' Sprinasjusl CA: Mono: Casa Di;oblo Min .. CA. SW ~. sec. 29. T to N. CA: Inyo: MI. Pinehol. CA. SW ~. sec. I. T I!S. 3S. R ) IE. 2104 ... 5_1.87. 80. Banner Spri,,&,. CA: R 33E. 1139 m. 4-1 7·87. SO. GoodaleCreek. CA: Inyo: Mono: Cua Diablo Min .. CA. SE \',!;«, 18. T 3S. R 1011. PilIcbol. CA. SE 'to sec. 17. T liS. R HE. 125 I 11 E. 2196 m . 5·1_17. 81. Spri", b~ TO"'cr Mine. CA: m. 4- 18·81. 51. Taboose CIftk. CA: In)"o: Bi8 PiM. Mono: Cna Diablo Min .. CA. SE \. sec. 3. T 3S. R CA. Sf v. sec. 8, T liS. R 34E. 1281 m. 4.18·87. 52. 31 E. 1952 m. 4.21.87. 82. Springs enlm", M"rt>k TinnnnahaCreek. CA: In)"o: Bis PIne. CA. NW V. sec. Creek. CA: Mono: While Mtn. Peak. CA. SE I. 5« 28. 28. T lOs. R 34E. 1281 m . 4.18·87. Sl. Sprinp N of T 2S. R 32E. 1525 m. 83. Marble Creek al HW 6. CA: Tinnemaha Creek and W of Poveny Hills. CA: Inyo: Mono: While Mtn. Pak. CA. NE ~. sec. 28. T 2S. R Bi& PiM. CA. NE 't. 5«. 30. T lOS. R J4E. 1403 m. 32 E. 1S15 m. 4·21_87. 84. Springs ca. 3.0 km S of 4-18·81.54. Bil Pine Creek. CA: Inyo: Bil Pine. CA. Benlon. W of HW 6. CA: Mono: Benlon. NV_CA. SE SE V. we. 24. T 9S. R B E. 1373 mA.18·87. 55. Baker ~'. sec. 8. T 2S. R 32E. 1617 m. 4.21_87. 85. Benlon Dftk. CA: Inyo: Bil Pine. CA. NE 't. 5«. 13. T 9S. R HOI Sprinas. CA: Mono: Gl,," MOllnl.ain. CA_NV. SW 3)E. 1112 m. 4.18·87. 56. Bis Pine Spring. CA: Inyo: 'f. 5«. 2. T 2S. R 31 E. 1708 m. 4_21_8 7. 86. Sprinas Bis Pine. CA. NE ~. 5«. 16, T 9S. R l)E. 1891 m. ~t N end of Blind Spri", Valle)". CA: Mono: GI,," 4·18·87.57. Spri",on hill SofWal"tnI Lake. CA: Inyo: Mountain. CA_NV. SW 't. 5«. 36. T IS. R ) IE. 1708 Bia Pine. CA NE >;. 5«. [6. T 9S. it H E, 1220 m. 58. m. 87. Sprinp,,1 Bramlctle RlInch. CA: Mono: Benlon. Springs in canyon W or W,,~n LHe. CA: lnyo: Big NV-CA. SW ~. 5«. 6 (springsnail posil;,",,). SE ~. 5«. Pine. CA. SW y. 5«. 2. T 9S. R llE. 1151 m . 4.18· 6 (ncp l i,"~). T IS. II. 12E, 1678 m. 4-21 -87. 88. Siream 87. 59. Sprinp in can)'on S of Shannon C"nyon. CA: in Long John Canyon. CA.: In)'o: Ne,,· York B IIII~. CA. IIIYO: Bi& Pine:. CA. SW h 5«. B. T 8S. R BE. 1373 14.4 km N_NE ofSW comer of quadrangle. 1769 m. m. 60. Sprinp N of Klondike We. CA: Inyo: BilPine. 5-4·87. 89. French Spring. CA: In)"o. Nc'" York BUlle. CA. SW V'i«. 23. T 8S. R 34E. 1208 m. 5·1·87. 61. CA. 8.8 km S of NW comer of quadrangle. 1617 m. FrtImWI CI"«k. CA: In)'o: Big Pine. CA. NE 't'5«. 90. Spring in WilIo'" Springs C""yon. CA: [nyo: In_ 20. T IS. R BE. 1434 m. 4-U·H. 62. Spring S of dependence. CA. ~.5 km NW ofSE comer ofquad· Ra_ Creek. CA: In)"o: Bishop. CA. ttl\ter of i«. rangle. 1861 m. 8_18_87. 9 1. Ba~1 Springs. CA: Inyo. 6. T IS. R BE. 1434 m. 4·18·87. 63. Spring in Chip­ Independcn«. CA. 13.6 km SW ofNE comer of quad­ munk Canyon. CA: In)'o: Bishop. CA. SW h i«. 21. ~ ngle. 1'152 m. 92. Sprina ca. 1.6 km SE of Mule T 7S. R 32E. 1708 m. 4·20·87. 64. Slream in Bulter· Spring. CA: In )"o: "'aucoN MIn .. CA. 11.7 km N-NE milk Cou.nl:l"y. CA: Inyo: MI. Tom. CA. NE 't, sec. 31. ofSW comer of quadrangle. 1586 m. 93. Milk Sprina. T 7S. R 3 1E. 2349 m, 5·1·17. 65. SI~am in McGee CA: Inyo. \\'aucobl Min .. CA. SE y, 5«. I. T lOS. R Meadow. CA: Inyo: MI. Tom. CA. NE V. sec. 22. T 34E. 1312 m. 94. Sprinlal Graham R ~nch. CA: In)·o. 7S. R liE. 1100 m. 4·24_87. 66. Spri", in DttpCan­ Wallcoba MIn .. CA. NE t~ sec. 12. T9S. R 34E. 1373 )'<>D. loll. Tom. CA. ~e n tn of K<. 12. T 7S. R l iE. Ill. 'IS. UI)IIlqCI Spron,. CA. In}·o. Wau. CA: Mono; MI . Morrison. CA. NE 'I. sec. 2.0 km S of Decllambcau Crttk along Hll> 395. CA: 6. T 4S. R 29 E. 2105 m. 4-29·87. 118. Springs al HOI Mono; Bodi~ . CA. SE 1'. s«. 30. T 2N, R 26E. 1922 Cretk Halchery. CA: Mono: MI. Morrison. CA, SW 'I. m. 5_6_87. 145. Mill Crttk. CA: Mono: Bodie, CA. sec. 35. T 3S. R 28 E, 2166 m. 4_ 23_87. 119. Springs NW 'I. sec. I S, T 2N. R 25 E, 2379 m. 5-~_87. 146. al HOI Cretk Ranch. CA: Mono; Ml. Morrison. CA. Slream N of Mill Creek. CA: Mono; Bodle. CA. SE '4 SE '" sec. 35, T 3S. R 28 E. 2166 m. 4-23-81. 120. Casa sec. I, T 2N. R 25 E. 2105 m, 5·6-87. 147. Stream in Diablo HOI Springs (cool seeps). CA: Mono; MI. Mor­ Rattlesnake Canyon. CA: Mono: Bodie. CA. NW 'I. rison. CA. NW 'to sec. 32. T 3S. R 28 E. 2257 m. 4·23- s«. 30. T 3N. R 26E. 2288 m. S-6-87. 14'. Murphy 87. 121. HOI Creek. CA: Mono. MI . Morrison, CA. Spring. CA: Mono: Bodi~ . CA. SE 1'. sec. 24. T 4N. R NE 'I. sec. 25. T 3S. R 28 E. 2196 m. 122. Layton 26 E. 2501 m. 5_6_87. 149. Siream S ofM ... rphy Spring. Springs ( ... nnamed on lopographic sheet). CA: Mono. CA: Mono: Bodie. CA. NE 'I. sec. 25. T 4N. R 26E, Casa Diablo Min .. CA. SE If. sec. 36, T 3S. R 29E. 2410 m. 5-6·87. ISO. Stream in COl1onwood Canyo!l. 2074 m. 123. Springs feeding l illie Alkali Lake. CA: CA: Mono: Bodie. CA, SE 'I. s«. 32. T 4N, R 27 E, Mono: Mt. Morrison. CA, SW II. sec. 20. T 2S. R 29E. 2227 m. 5-6-87. 15 I. S~ l ph ... r Pond and pool 10 E. CA; 2104 m, 4-29·87. 12 4. l illie HOI Crttk (so ... rce). CA Mono: Trench Canyon. CA-NY. SE If. sec. 26, T IN, Mono: M1. Morrison. CA. NW If. sec. 13. T 3S. R 28E. R 27E, 1952 m. 5-6·87. 152. Saline Marsh. CA: Inyo; 1074 m. 5-6-87. 125. Deadman Creek. CA: Mono: MI. Ne ... York BUlle. CA. eenlCT of sec. 27, T 14S, R 28E. Morrison, CA, NE 'I. sec. 27, T2S, R 27E. 2257 m, 335 m. 153. Sueam in H ... nlerCanyon. CA: In)'o; New 4-29·87.126. BigSprings. CA: Mono; Cowuack Mm.. York B~ll e, CA. 14.5 km SE of NW comer of quad­ CA. NE 'to sec. 25. T 2S. R 27E. 2166 m. 4_ 23_ 87. 127. rancle, 549 m. 3-28·87. IS4. Siream in Bevaid&cCan­ Spring ca. 0.5 km E-NE of Arculari .... Ranch. CA: Mono; yon. CA: Inyo: New York BUlle. CA. 12.3 lim E-SE of Co,",uack MIn .. CA. SE 1'. sec. 20, T 2S. R 28E. 2 135 NW carner of quadrangle, 580 m. 1_28·87. 155. Bad· m. 5-6-87. 128. Springs along lowcr seclion of Mc· waler Sprio8- CA: Inyo: Waucoba Wasil . CA, 10.0 kin Laughlin Creek:. CA: Mono; COWlrack MIn .. CA. NE NE ofSW comer ofquadranglc. 488 m, 3-28-87. 156. 'I. sec. I S, T 2S. R 28E. 2318 m. 5-6-87. 129. Mc_ Sprinp S of Lower Warm Sprinp. CA: Inyo. WallCOba Laughlin Spring. CA: Mono: COWlrack MIn .. CA. NE Wa$h. CA, NE 'to sec. 30, T 13S, R 39E. 381 m. 3_29_ 'I. sec. 12. T 2S. R 28E. 2654 m. 5-6-87. 130. Springs 81.157. Springs NW of Upper Warm Springs. CA: 31 S end of Black Lake. CA: Mono; Glass Mountain. In),o; Waucoba Wash. CA. SE '" sec. 5. T Us, R 39E. CA-NY. SW 'I. s«. 4. T 2S, R 31E, 1922 m. 4·21_87. 549 m. 3·29·87. 158. Upper Warm Spring. CA: loyo; 131. Sprin8" al N end of Black Lake. CA: Mono; Glass Dry Mountain. CA. NE 'I. s«. 9. T 13S. R 39E. 488 Mountain, CA-NV, SE 'I. sec. 29. T IS. R 31E. 1952 m. 2- 11 -85. 159. Willow Crttk. CA: Inyo; Waucoba m, 4_21·87. 132. Ri ver Spring. CA: Mono. Glass Wash. CA. 12.3 km N·NE ofSW corner ofquadranak. M o~n'ain. CA_NY. NE '1.~ . 24. TIN. R JOE. 198J 915 m, 3-2i-i7. 160. COITIII Sprinso· CA: Iny<>; BlaD«> VOLUME ]02. NUMBER]

Mtn .. CA, SW If, see. 3. T 8S. R 36E. 1525 m. 16 1. CA. 11 .3 km SW orNE comer of quadrangle. IJ73 m. Bot. M01,U1ds Springs. CA: Inyo; Blanco Mtn .. CA. SW 2.IS.8S. 189. Seepage on SW side of Panaminl Lake \Iosee. 32. T 7S. R 36E.] 508 m. 4·26-87. 162. Antelope bed. CA: Inyo: Manlcy Peak. CA. E sides of sccs. 28 Spring. CA: Inyo; Blanco Mountain. CA. NW If, see. and 34. T 22S. R .wE. 317 m. 3·31-87.190. Slream 24. T 7S. R 35E. 1708 m. 163. Samms Spring. CA: in Pleasant Canyo~. CA: !nyo: Telescope Peak. CA. inyo; Blanco M01,U1lain. CA. SW If. see. 13. T 1S. R 6.9 km NE 01 SW comer 0 1 quadrangle. PH m. t9\. JSE. 1739 m. 3·29-87. 164. Willow Spring. CA: I n~o: Past Offitt Spring. CA: Inyo; Telescope Peak. CA. NE Mapune. CA. 10.6 km SW of NE corntr of CA. 15.3 km NE ofSw com er ofquadransle. 549 m. quadrangle. 366 m. 3-14·85. 215. Warm Spring Ck ).)1_87. 188. Anvil Spring. CA: Inyo: Manley Peak. In)'o; Wingate Wash. CA. 7.2 km SE ofNW comer of '" PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON quadran&lc. 7)2 m. 2-18-8S. 2 16. Lower Talc Mine 2,8 km NE ofSW comer of Quadrangle. - 79 m. 2)2. Spring. CA: 111)'0; Bellneus Well. CA. 6.4 km £ _NE of Salt Springs. W ofCouonbali Marsh. CA: I ~yo: Chlo­ SW romerofquadrangle. 9lS m. 3·10-87. 217. Willow ride Cliff. CA·NV. 1.0 km NE of SW cornu of quad­ Spring. CA: 111)'0: Funeral Pt-ak . CA. 7.8 km N E of SW ran.gle. - 73 m. 2-22-85. 233. Salt Springs S of Cow comer of quadrangle. 732 m. 3-20-87. 218. Hidden Creek. CA: Inyo: Furnace C=k. CA. SE ... !lee. 3. T Spring. CA Inyo: Funeral Peak. CA. 1~.3 km NE of 27N. R [E. 31 m . 2_5.83. 234. Stream in Omonwood SW romer()fquadran&le. [)42 m. 3-]7-87. 219. Springs Canyon. CA: Inyo: Marble Can)·on. CA. stream 10.0 al Mormon Point CA: /11)"0; Benneus Well, CA. 6.7 km NW ofSE comer of quadran&le. 732 m. 2·iS-8S. km N_NW of Sf comer of quadrangle. -64 m. 3-9- stream 11.2 km E-NE of SW comer of QII8dranaIc. 85. 220. Spring (Io""r) in Johnson Canyon. CA: In yo: 1037 m. 3- 12-87. 235. Mclean Spring (Salt Creek). Telesrope Peak. CA. SW 'A sec. 22. T 2 1S. R 46E. 1190 CA: Inyo; Stovepipe Wells. CA. SW 'I. sec. 31. T ISS. m. 221. Ea$.le Borax Spring. CA: Inyo; Bennells Well. R 46E. 37 m. ).8-8S. 236, Springs NW of lL""3ne WOJI ­ CA. 13.3 km SE ofNW comer of quadrangle. 79 m. der Mine. CA: Inyo: Ch[oride Cliff. CA-NV, SE 1'. !Iee. 2-16-85. 2n. Hana upal\ Spring. CA: Inyo: Telescope: I. T [5S. R 46E. 366 m. 2- 13-85. 237. Monarch Sprin,. Peak, CA. 1.6 km S_Sw of NE comer of quadran&lc, CA: Inyo; Chloride Cliff. CA-NV. SE 'I. $«. 24. T 14S. 1281 m. 223. Bad""aler Spring. CA: 111)"0; BennC'lI$ R 46E. 915 m. 2-4-85. 238. Keane Sprins. CA: Inyo: Well. CA, 2.7 km SW on NE comer of quadrangle , Chloride Cliff. CA_NV. NW 'I, sec. 8. T JOW, R IE. - 79 m. 2-16 ·8~ . 224. Tule Spring. CA: Inyo: Bennetts 1159 m. 2-4-85. 239. Cane Spring. NV: Nye: Cane Well. CA. 10.8 km E-SE of NW comer of Quadrangle. Spring, NV. 6.0 km N-NEofSW eomerofquadran&k. - 79 m. 2-28-85. 225. Grapevine Springs. NV: Nye; ID8 m. 240. Spring in Oasis Valley, 13.2 km N of Ash Meadows. NV·CA. NE 'I, sec. 2. T 19S. R 50E. Beauy. NV; Nye County: Thirsty Canyon. NV. SE 1'. 732 m. 3- 14-85. 226. Blackwater Spring. CA: In),o; $«.32. T lOS. R 47E. 11 28 m. 241. G rapevineSprinp. Emigrant Canyon. CA. 12.9 km S-SW orNE com er of CA: Inyo; Ubehebe Crater, CA -NV. NE "'!Iee. 3. T Quadrangle. 946 m, 3_10_87. 227. Navel Spring. CA: liS. R 42E, 824 m. 242. Spri~ 1.6 km EofSrotty's [nyo; Ryan, CA-NV. NW 'I, sec. 13. T 26N, R 2E. 640 Castle. CA: Inyo: Ubehebe Crater. CA-NV. NW 'I. $« m. 3-17-87 228. T ra" enine Springs. CA: [n)-o; fur­ 5, T 115. R 42E. 946 m. 243, lillie Sand $prill&- CA: nace Creek. CA. NW 'I, sec. 25. T 27N. R IE. 122 m. [n)'O; LaSt Chance Range. CA-NV. SE II> $«. 17. T 95, 229. Salt Springs. NW of Cow Creek, CA: Inyo: Chlo· R 41 E. 915 m. 387. 244. Sand Spring. CA: Inyo: Us! ride Cliff, CA_NV. SW 'I, sec. 21. T l8N. R IE. -79 Chantt Ran&,!. CA_NV. SE II> sec. 7. T 9S, R 41E. 946 m, 2-J-85. DO. Nevare. Spri~ •. CA; In yo; ChlQridc on, 3-30-87. 245. La.. C h an~c Sprin" CA: Inyo; Ma­ Cliff: CA-NV. NE 'I, sec. 36. T 28N. R IE, 275 m. 231. gruder Mtn" NV-CA, SE 1'. sec. 2. T 8S. R 39 E. 1739 COllon bal1 Marsh. CA: In}'o: Chloride Cliff. CA·NV. m.3-30-87.