Arizona Hydrobiidae (Prosobranchia: Rissoacsa)

Robert Hershier

and J. Jerry Landye ABSTRACT

- - Hr..7h1P-r, Robert. and J. Jerry Landye. Arizona Hydrobii ,

(Frosobranchia: Rissoacea).-A diverse fauna of Arizona

Hydrobiidae IS doritmentr-d as a result of recent collecting from

numerous springs in the state. The fauna is comPosed of 14

smecies in two g.=ner,... , Evrmulomr4i= CA.11 and Filsbrv and Trvonia 4 .4.-_ 5tiMPSOM. ,n r =mxciPs are described as new nerein arc tne

fourteenth represents a new record for the state. All of thi=

sPecies have relatively rr-.=.trir- ted distributors. and two are

SINSLE sPring endemics. All congeners are allcmatrir- to one

anr,ther.

StaPwise canonical discriminant function analyses using sets of

and anatomical data confirmed the distinctiveness of the 12 species' of Arizona FyrguloPsis. as classification of (grlip,=.d)

topotypes was 89-93% based on analyses using shell data. and 100%

for analyses using anatomical data. The discriminant analyses also confirmed that diff nti...-ttir,n ,amonp congeners largely

involves male anatomy (especially menial features), as this data set yielded the best separatirn of species in plots of scores on

the first two discriminant functions.

2 Robert Hershler, Department of Invertebrate Zoology, National

Museum of Natural History. The Smithsonian Institution,

Washington, D.C. 20560. J. Jerry Landye, Arizona Game and Fish

DePartment, HC 66. Box 2205, Cornvills. AZ 66325.

3 Arizona ;yOrotiidae (Fri sobranchia: Rissoacea

Robert Hershier and J. j:=rr.,,, -=1 .ndv.=,

Introduction

caobranch sn: i of the -F asnil , Hvdrobiidae comPrise a diverse 3 4444411. id to71)414411L-Via component of the .gr..-44.1i,=;..4.- -F auna (di the American with A numerous locally ehd:=171i7 ieoleted complexee or drainage te!:1- t'T-J- d-u=nout tn-E r;A.4 1966; Land 1977 in P rt H11.= t71 4.4.0:- 0 - colisctr and studying thess minute nei la. th,=,v remain Poorly known in the 44--- VeT4P- N . - , The:-e are no publienedA mods:n. r7 nem=ivi= - - 1441.4.0 systemat c tratmen o " -f aunas - - but ass A

Taylor, 1987Aand what little work has teen done larei 0=Si-57E of isolat;.d ecies descriPtions based cn study emt shell. There is a clear and urgent need to address the systematics of this fauna given the f=(f- t that many .B.4-44=b4i.ci.e

OAR, , - . r=c- r LT.-rcl;..t..14: threatened or endA.ngerd as a r,==,,Itit of A deterioration or disaPpearance of natural aquatic habitats 4.A.-44-gra.

(Taylor, 137('a). As an illustration, 12 of 21 southwestern species being considered for federal listing threatened or endangeredcre und;=srribedie(lai., 1984) A To further this end, we present ir a systematic description of Arizona Hydrobiidae. To date, this fauna has been largely ignored by malacologist=.. Prior to 197, the __le published record of live-collected hydrobiids from the state was by Pilsbry and Ferriss (1909)A who obtained Paludestrina stearnsiana from a i)v single spring in the Huachuca Mountains. In their monograph on

4 Arizona mollusks, ;;;...guart and Miller (1973) listed EiX additional such record far the family. rePresentinp sP.vPr,:kl possible species. Additional records of collections of Recent or subfossil shell are listed Taylor (1970b, 1975).

In the past 20 years the junior authar has intensively surveyed permanent Arizona waters and found 4.0reol a lar:pe hydrobiid fauna)

with P.,:timatx.ri SCCI1C divf=rsity raning upwards to

18 sPecies (see Landv,.. 1973; Landve. 1581; William= ea al..

192E). The bulk of thi=, coll Ection. larPe alcohol holdinpE from

localities in the state and PeriPheral areas ;Fig. 1). WSE made available to the senior author- whose taxonomic work is

.- - pr.=---,=ni-:,=d herein. rt .=.,=p.....r:A.t-,= p.,-4.pr tr .-.-4.ting .. ologic=.1 :,nri biopeograpical aspects of the fauna will be published elsewhere.

-c Fourteen sPecies are recognized 47; t.7 iii 4--'r representing two -.> genera, Tryonia Stimpson, 1865 and Fyrpulopsis Call and PilEbry,

1886. Thirteen of the species are new. and the fourteenth. SEA& PyrPulapsis deserta (Pilsbry). represents a new record. A _,■0=' tot S tepwise discriminant function analyses using separate

SHELL and anatomical data sets were performed in order to examine

-, - , PAT TERN A of variation and tR.= t tql 0=th:=r the 12 new. alloPatric

FyrpuloPsis app. are distinct -From one another.

F 1.1'7,T OF RED:GNU:ED SPECIES

Fyrgulomsis glandulosus Hershler, n. sm.

Fvrgulomsis deserts (Pilsbry)

Fvrquiomsis bac:hus Hersrier. n. SP.

P'yrpulomsis conicus Hersrler, n. sm.

Fvrpulopsis morrisori Hersrier, n.sm.

PvrpuloPsis mon1e7 srsHs Hsrshlsr, m.

P',rg loPsis sol_a Hershier, n. sp.

F\irmuloPsis simPlex Hersrier, n. sp.

Fvrgulomsis sancarlr,sersIs Hershler, r. sp.

Pyrmulomsis cflrHluentis H fliCr fl. Esp

Fvrmulomsis cocrisi Her-shier. n. SP.

FYr7U1OPSiS CMPSC:i Hersrler, n. sm.

Trycnia milae Hershisr, n. SP.

Tryonia guitobamuitae Hershler, n. sp.

MATERIALS AND METHODS

MATERIAL EXAMINED. Material from 35 localities (Fig. 1),

totalling 48 lots, was collected uci/114 a fine-meshed hand sieve,

relaxed with menthol crystals, fixed in dilute formalin and

preserved in 70% ethanol. Most eii.40felre lots --- t erreht.- 100 specimens; Aocality data -ibt-i4a—tizsdabzia•-e7:..*--e-i-tr.F-A- ." are in Al! Appendix 1. Th.s. cf this material is housed in the

United States National Museum collection (USNM catalog numbers ma, are referred to below), ;Ether relevant material from thOlar

collection and that of the Academy of Natural Sciences of

6 Philadelphia ( ANSR) was also examined.

MORPHOLOGIC STUDY. Shells and animals were studied and

measured at 50X using WILD M-5 and M-7 dissecting microscopes

egitipged with ocular micrometers. Shells. ogerculae and radulae

were photographed using a HITACHI 5-570 scanning electron esem) micrcscoPe. Ts obtain intact bodies for disssctisn 4 shells were A de.dalciiied by scakinT in ccncentrataz Eduin = .A.P.4.4404,4" th ,= r=maininq pe=1 1-idle was AMiM:4, 1:7" were di===,... ,,,Jd in

dilute Ebuin's Sclittif7n. 7c examine penial mor7,holo.=%/ and aA44.444A ciliation patt ,=rn= on tn= neni and

w=re dried g=inq nr-7NTTIN DCF-1 Critical Foint Drier 4f.pf tni=m

•

C ,7ntinurots and ( to a lesser extent maristiz data were obtained 544,erid 71. 4 1/ from hei 1 and anatomy sgesimens

Pvrgitlogsi= ( Table 1) for use in d scriminant

function analyses.

______She 1 La were

cleaned with CLOROX to remove surface dePcsits) .F4102.001i44

41=0Me*w*Or5+ oriented in the standard aPertural

aspect. 4strweca44.1wavaliwuket(werse am14.a... first pinPointe477-777;77a0

necessary points 44..em or aspects of 44.4., shell outlin:r:nto paper

using a camera lucida. Distances between 411: aQ. points were then

determined using a fine-scaled millimeter ruler. The following

characters were measured or counted:

(1) number of whorls (NW)

(2) shell hei9ht (SW

7 (3) shell width (SW)

(4) aperture heiciht (AH)

(5) aperture width (AW)

(6) height of bddy whorl (LEW)

(7) width of body whorl (WELL)

(I;) 1,=ngtn of line ronn=cting sutures at posterior ends o+ body and Penultimate whorls

(9) maximum Pe!-- gendiduiar distance between line joining the a 7vc= =utur S. and outline c+ penultimate whorl

The +allowing ratios ware -F ormed UEiMq the above data:

(1) =n=.11 height _shell width (ZH/EW

nc ht (:::( H/EH)

(3) body whorl lenqth/sh= 1 1 height (L7M/SH)

(4) whorl convexity (CD/ AE), +allowing measure provided n and Riggs (1975: F. 352)

Anatomical counts and measurements were taken from five males

4416‘ and five females from lot. Body and gonad lengths were

6 measured with the animal pinned iiihmost in 5 41.. apical aspect. The penis was cut from the neck. and 40,A measured in 4.19,e- dorsal aspect. The pallial roof and visceral coil were removed from the head/foot and 4400 separated from one another along the end of the pallial cavity. Fallial structures were measured with the pallial roof flattened eftm4 and pinned. The following counts and measurements were made (1-7, both sexes; 8-21, females: 23-30, males):

(1) body length (EL) (2) oePhrabium length (CI L)

(3) osPhradium width (DEW)

(4) distance from posterior end of osphradium to Posterior tip of ctenidium (OCT)

(';) ctenidium length (CTL)

(6) ctenitium width (1-7W)

7 C, number of ctenidial i'laments

(8) ovary ienqr (CL.

(9) caPsule l no lenczth (Cf;L)

(1()) capsuis glanc widtn G2GW)

1 - ,7 ( 11 ;,11.;UM=7 p1 -Ino lemsrh ( CL)

(12) albumen sland width (Jf Eli,;

(13) diameter (anterior-postrior) anterior coil of oviduct

(EC' )

(14) distance from anterior End of oviduct coil to posterior end of capsule gland (COP)

(15) maximum diameter of anterior portion of oviduct (COD)

(16) bursa copulatrix length (BUL)

(17) bursa copulatrix width (KM)

(18) width of bursa cogulatrix duct (BDW)

(19) distance from posterior end of albumen gland to posterior tip of bursa (BAG)

(20) seminal receptacle (body plus duct) length ( SRL)

(21) seminal receptacle width (SRW)

(22) testis length (TS)

(23) prostate gland length (LPR)

(24) prostate gland width (WRR)

5 (25) 1em3th of Fenis from ditse to tip Fenial lobe (L2)

(26) Penis width (at base) (EW)

(27) distance from base of penis to b..=,.se of Penial filament (L3)

(28) Fenial filament lerczth ( LFI)

(29) penial filament width (WFI)

(30) number of glandular r1d9es on penis (GLR)

- , - -, Ti s followin7t fi7rmr.-d it ing these amatomii t1

(i both sexes: 4-. females: 10-14. males):

(1) osphradium 1F-TIgt, 1 enidium lenth (DELSCTL)

(2) distance from posterior tip of osphradium to ctenitium t iF 1A,ngth (OCT 'CTL)

(3) ctenitium 1 1-1qth (171-1 )

(4) ovary length/body 1=,ngth (n)/7,1 )

(5) caPsule gland length/albumen gland length (CGL/AGL)

(6) oviduct coil diameter/albumen gland length ( COL/AGL)

(7) bursa length/albumen gland length ( BUL/AGL)

(8) distance from albumen gland tip to bursa tip/bursa length

(BAG/BUL)

(9) seminal receptacle length/bursa length (SRL/BUL)

(10) testis length/body length ( TS/BL)

(11) prostate length/body length (LPR/BL)

(12) distance from base of penis to base of penial filament/penis length ( L3/L2)

(13) length of filament/penis length ( LFI/L2)

(14) filament length/width ( WFI/LFI)

10 The total character breakdown is as follows: male and femzie shell, 5 raw characters, 4 ratiost(totallinq 13 character): male 0°4 4 anatomy, 16,1 8f-.1 (24; female anatomy, 21/efraA4. .

DATA ANALYSIS. DescriPtive statistics were generated, and analyses of variance (ANOVA), t-tests, and stepwise discriminant

, . function analyses were performed using the c'FS.-X program. subPrograms BREAKDCWN. T-TEST. and DI7,17RIMINANT.

47 1.... MAHAL rxtr-LIPWi'= - procedure W&E used in C3Z5 discriminant analyses, in which vAri..11.bl4=F are selected on the basis of

- providing the largest MahalAnobi= distance between the cln=;-=t pair of groups. $7:EPWiSE procedure was usedas it s! lows rPrognition of the most effectively discriminating variables.

- - e04404i1UAd4 Topotypes of er h =.p.F.ciss were is.1;pa ed as 12 s.=parate ,lroups, with other populations inputted as unprour-"ede ernm.• to prevent biases resulting from having data available for varying IA 01.4M , numbers (one-four) of populations for the 0 spe-ig.=.. All populations. used Ripm■-444.peo--ewie are listed in Table 1.

Separate analyses (including raw plus ratio data) were Performed not only for male and female anatomical data sets, but also for 44,-414C.444 male and female shell data, as =..- xual dimorphism in size

in at least one population -For 10 of 12 species and in 11 of 23

total populations studied (Appendix 2). The pooled within-qroups covariance matrix' was used to comPute probabilities of proup

memberehip.

11 AnKNOWLEDEMENTE

We thank Drs. G.N. Davis and W. Fratt -Frlr lending material.

Paul Greenhall measured hel and prr-cvid;=d other diverse technical assistance. Dr. Lm.-Ann Have :: helped gratly with lni retat Ion 01' data an.4.1y=rz-=. and Ms. Cindy Carman and Mr.

, Faipn iThapm,in ran the ..=t pproPriate cdmPuter programs. Nolly Ryan prP.-=.. ared mdst c.f the .final illustrations and Plates. G.M.

Davis. F.G. ThcmPson and W.F. Fonder provided use-Ful criticism of the manuscriPt.

Hershier's particiPaticn in the prcject was -F undeo, in Part, ty r 2 O'rt Sikh (),v/v 7 Fart

, -f rr m thr7,ugh 1 '1771 by grant= to Dr.

W. -. L. -J a-444 tAtr-Ct Fi=h =knri e L44.. Z.4a 4 Servicim

SYSTEMATICS

Accounts for each species consist o+,4#he—P RZ provision of common name; synonomy: material examined: diagnosis: description; remarks (where applicable); type locality; distribution; and etymology (where applicable). Designation oi common names 4a44.--t.V.0.e-enetv-t-TinT7- largely follows that of Landye

(1781) and Williams et al. (1785). host ei--449.s. species 464,44

1444riTi9ItT^,TP were recognized in unpublished reports by Landye (1773,

158 1 ). Our synonomies include references to these works only

12 when 44441. identities of his taxa are unclear. Congeners

hi="=', are net greatly divergent in morPhology, descrimtions ARE 4r-pe+ae-fLetaf..- provided for only one species Per genus, with other descriPtions emPhasizing distinctive f,=,..e turP=-

Such features include those (in order of importance) involving penial morPhology, shell, head-foot and penial pigmentation• and morphology of thx b!trsa ccoultrix czmplEx.

- . R.SZKDULAR DATA ARE IN LABIA 2. SH ,,--. 11 AND .=I NATDMICAI DATA

FYRULOPSIS SpP. ARE in Ap=sndideE 2-4.

Faml HYDRFL 7.-II2AE

and F lstry. lEE=

REMARKS. Adeai led cen ic DR=S7RIPTICN ( WITH syncndmias! is provided by Hershler and Thompson (in press), in which Marstonia

F.C. Baker, 1926; Fontelicella Gregg and Taylor, 1965;

Microamnicola Gregg and Taylor. 1965; Natricola Gregg and Taylor,

1965; and Mexistiobia Hershler, 1585Aare considered synonomous with Fyrgulhp,,i=. PyrgulePsis, as presently understood, have globose to elengate-conic shells and penes with a small lobe, narrow filament, and one to fifteen glandular ridges, sometimes on stalked crests. The genus occurs in much of eastern North

America as well as in the Southwest and northern Mexico and comprises 35 (including the below) described species as well an additional 20-25 undescribed species from the Southwest. The closest relative of the genus appears to be Cincinnatia Pilsbry,

1891, which is restricted to eastern North America.

13 PyrguloPsis glandulosus Hershier, new sPecies

Verde Rim sPringsnail

Figs. 2a-c, 3-E, 9a, b

FONTELICALLA sig. Williams et al., 1985: 19.

MATERIAL EXAMINED. ARIZONA. YAVAPAI COUNTY: Nelson Place Egring, USNN 8:3047 (holntype), U5NM E59048 (5 paratyPes), USNN E47205. J.J. Landve and G. Edward=. 25 Si=Ptember 1'473: SPring 150 m E oi Nelson Place Spring, H5NN F..k 47232, J.3. Landye and 3. Edwards, 227 SeptembPr 1577.

DIAGNOSIS. An ovate-shell=d. large-sized sPecii==. elongate and massive., filament relatively small. Dorsal genial

=grf?ce with several glandular ridges, two or more of which are elongate: ventral =Atrfan,= with additional ridges borne on

,- accessory rc-==t=. Holc zpe (Fig. 2a) height, 2.80 mm; width.

1.92 mm; whorls 4.0.

DESCRIPTION. Shell whit=,-ciear, tr..- n arant; periostracum light brown, thin, covering much of shell surface. or absent.

Shell (Fig. 2a-c) about third than wide: height, 2.0-2.S mm. Whorls, 3.5-4.0, convex and slightly shouldered. =gtur;.“= slightly impressed. Spire outline convex, largely due to rapid translation of body whorl along shell axis. Sexual dimorphism significant (Appendix 2) with females larger than males.

Protcconch slightly protruding, with 1.25 partly pitted whorls.

Body whorl relatively inflated and large (8374 of shell height).

Aperture ovate, moderate in size. occupying about half of shell height and E3% of body whorl heieht. somewhat angled abcve rounded below. and tvpic=lly loosened -from body whorl. Inner lip slightly thickened, moderately reflected and fairly =tr.Right.

14 Outer li thin and tro4.Oly co 7: Prtur5l Plane near. i ...-

PARALLEL , TO COILING AXIS WITH LONG Rxi=. TILTED COG R I== .77

RELATIVE TO COILING AXIS. UMBILICUS MODERATELY to BROADLY OMEN.

Teleoconch SCULPTURE CONSISTING OF STRONG GROWTH LINES. Amber.

- - horny, PAUCISPIRAL OP.=RCULUM ABOUT c71 0= AND A HALF TIMES AS LONQ

AS wi P. and WITH THREE WHORLS.

, - 7NOUT than wir"e. im.irly thickened =1.n ,-i terminat D1= 7:E:11 ;,,q17h 1:1G-F.ny CR.S.1 l=pnPt=. CPphalio tentacles (Tn) narrow. sii=ntly less .1: -; an twice as long as snout. , and somewhat expariciPt at tiPP. IrrP 17t r t[tit= O cilia on tentacle surface ;.=in:J. 4b, cf. =am=tim=s for-m.1mq a short ti-act en laft tentacle. BiaL located IN SLIGHTLY SWOLLEN

EDGES TEN TSC OASES. ELONGATE FOOT TNIC*SHED.

POSTERIORLY TAPERED. ANTERIOR EDGE OF FOOT FAIRLY

WITH CENTRAL PORE THROUGH NCH TWELVE TO FIFTEEN elongdte EAL

GLANDS DISCHARGE.

EPITHELIAL MELANIC piiilment usually caeR044oadbiar ABSENT FROM head/foot ALTHOUGH OCCASIONAL SNAILS MAY HAVE LIGHT TO FAIRLY

DARK BROWN PATCHES COVERING MOST OF SNOUT (POSTERIOR TO ORAL

LAPPETS). MELANIC PIGMENT COVERS much OF DORSAL SURFACR, OF

VISCERAL COIL (FIG. 3), WITH STOMACH REGION TYPICALLY SOMEWHAT

LESS PIGMENTED OR EVEN LACKING PIGMENT. LIGHT GREY TO DARK BLACK

SUBEPITHELIAL PIGMENT sometimes OCCURRING AS SMALL PATCHES JUST

DISTAL TO BASES OF TENTACLES AND ON "NECK" BEHIND TENTACLES, AS A

BROAD TRIANGULAR PATCH ON SIDES OF HEAD/FOOT. AND AS AN IRREGULAR

PATCH ON OPERCULIGEROUS LOBE. SIMILAR PIGMENT PRESENT ON right

SIDE OF MANTLE COLLAR. OPAQUE WHITE GRANULES DENSELY CLUSTERED

JUST BEHIND EYESPOTS, WITH SMALL CLUSTERS SOMETIMES OCCURRING

4 5 along the proximal half of tentacles as well. Dark, brown-black

subepithelial pigment concentrated in proximal portion o-F. penial

filament, extending somewhat onto rest of penis (Fig. 3). While

visible when penis viewed dor-=Ally (as in Fig. pigment

actually positioned closer to ventral penial sun-ace.

rIrq.-7..nization shown in Fig. 5a.b. CtniHiltm

large. with 1 -Zf.") narrow filaments. filling most pallial

cavity length 24-23% of nori.y 1,=,ngth), and extending

anteriorly to mantle collar (Mc). Widest filaments centrally

, , Pt7=-7itinnri and hr t=ning acainat int =stine In aspnraCiitm

fairly elongate. filling 22-=. of ctanidium length. pr7-7itinrif=d

slightly pc snlor to Kidney with

small opening fringed by distinctive liPs.

Buccal mass filling anterior half of snout. Pair tubular salivary glands dorsal to nerve ring. Cesophag l tube simple

and narrow. Pair o+ chitinous jaws positioned at anterior end of

radula. Radula (Fig. 6) typically taenioglottg,, with

trapezoidal central tooth having single pair of basal CUSPS.

Lateral angles central tooth well developed, projecting about to 45 from central axis of tooth. Basal process of central tooth

well-developed. Central and lateral teeth with enlarged central

cusps. Outer and inner marginals similar in shape, with the

former having only a few more cusps than the latter. Stomach

(Fig. 5c, Ast, Pst) and style sac (Sts) about equal in length,

the former having a small caecal chamber (Cc) along its posterior

edge. Posterior stomach chamber with single opening to digestive

gland (Dgo) (posterior to oesophageal opening). Digestive gland

16 covering posterior half of stomach. and extending (ventral to gonad) almost to posterior tip of visceral coil. IntestinP (In) with slight bend in pallial cavity (Fig. 5b).

Testis (Fig. 5d) consisting of =iMPle lobes, with anterior end lying against posterior erigP of =-trmach. and filling about thrPP- quarters, of digestive gland length and 7fl%- of body 1Ph.-, th.

. Eemin.=.1v icle F . 5t) exit ..-tntr=ri71r end testis and consisting of a -Few thickened coils ventral to tsstie and partly covarin,= dorsal stomach surface. F ostate !-7 1and

, simPle, small (1&:%) i-'eiatiie to Ondv Pfl=t2rif7r to

(interior from mid-line o+

Prostate ant extending alon,:: cclumelier i ti side of aIliai

-,- , 7avity be-ors L11tf=rinc: "n=r- k" PrFt,=rinr to oar isi .., ttrhm=nt

(Fig. 7a).

Penis (Fig=. 7, 4, 5;4, 7, P.) 1..-irgP, protruding well anterior to mantle collar (Fig. 4a). flattened, broadly attached to neck well

PC sterior to snout (Fig. 5). Penis extending anteriorly from attachment area without cnilinp (Fig. 5A., Pn). Penis much longer than wide, often expanding in width distal to base. Penial filament (Fig. 7, Ff.) one quarter of penis length, positioned at about two-thirds of penis length, narrow (length/width, 25%), and rarely projecting distal to penial lobe (Fig. 7, Flo), which

tapers and sometimes has a distinctive terminal process (Fig. 7b. d). Vas deferens extending as fairly straight tube along edge of

Penis (Fig. 7a), and opening at tip of penial filament. 40 Penial surface with four-A-seven slightly elevated glandular

ridges, consisting of paired rows of small glands discharging through a central slit. Dorsal penal surface with two elongate

17

RI DGES. one positioned 1=trgP.ly along edge containing the ;menial filAment, and other located along opposite edge, sometimes curving toward the filament (Figs. 7 B. One to three additional and smaller ridgr=-= on same surface. typically

Pcsitioned on i7i0 ni pg.ni=sl lobe (vi=ible from w.ntrA.I =1=Pct also) and h=.tw=.f.=n two =long=tt= rlices. Ventral pamial sur-Fsrs with one to three rioqes lccatat on two Ç - - Crest

- loc-q.t7=d closest to distal ede of peni = somewhat swollen and bulging -from sniai surface whereas Œth5r or=.=t..7 tv7i.-- =d1v as A.C= 4 r= 4-

Ovary (hot l' iqursdY lobular. stuttinc anterloriv sTainst posterior (=OP:a stomach, anri f±11 Inc 17% of body lengtn.

Ovid:tct pass In over stomach =itr ace before disapPearing

(to left of) pallial oviduct. Left lateral Palliai oviduct and bursa copulatrix complex shown in Fig. 9a.b. Albumen gland (Ag) highly glandular and somewhat (104%) longer than capsule gland (CEO, which has Posterior. yellow section and

(smaller) anterior, clear section. Note that gland has a moderate pallial component. Capsule gland opening (Coat) subterminal and fairly long and wide. Anterior to openinq, presumed pathway of egg capsules fringed by thin extension of tissue from capsule gland (Fig. 9a), as typical of genus.

Oviduct (0v) with single loop proximal to point of merger with

. duct of seminal receptacle (Sr). Distal to this, oviduct merges with duct of bursa copulatrix to jointly enter albumen gland

(Ag). Simple oviduct loop small, with diameter about 44% of albumen gland length. Body of small seminal receptacle,

18 Pnsitinned en 1.=ft surface of bursa, often narrow and

indistinguishable from its duct; lenclth of body plus duct almost equal (837.) to bursa length. Bur. (Fig. 5b) moderately lar

(43% of albumen gland length), g=tr-,.... hained, and lying against

left side (near ventral edge) of albumen gland. Two thirds of

1=ngth posterior to gird. Thin narrow duct of bursa

tightly pressed to albumen gland.

REMARKS. PvrguloPsis glandulosus groups with a numcer ni

Arizona and New Mexico ePecies (thA., latter undescribed. see

Taylor, 19= having 1,-4.rge, ovate 5hr=11.- and I-7P4M with small . filament and relatively numerous glandular ridges. Cn the basis

penial merPhology, th.= =ni=cif.== i ,T4 meat aimiiar =. 411X mentezumensis. which also occurs in Verde River drainage. A TYPE LOCALITY. Nelson Place Spring, Yavapai County!

DISTRIBUTION. Nelson Fla.ce Spring complex. consisting of two

=prings separated by 150 m.

, ETYMOLCriY. From La tin glA.ndi =. referring to the highly

n ature of the penis in this species. Pvrquir.7-,sie Ceserta (Fil= ry)

Figs. 7c, 10a-d, 11,a-c, 13

Amnicola deae-ta Filsbr,. 1716: 111-112. figs E. 7.

N. gen., n. Lard,e. 1=777: 4E.

"Hmnicolan de rt. rdve. lcEi: 70-71.

MATEFIAI U:-H. WA9HTNizTC-IN 1701INTY: sT=.rin,72 on Old Ea'4tion Ranch, HEH1 7477f. 3. J. 1 -indye. 21 February 1977: =m,...,11 sPring, 70 m N c-f r i iF, 119NN 47247, 1,J. L.=..ndyF4. 21 February 1'977: Green St:ring. USNM 647242. j.J. Landye, - February 1977: =7:-,:'ing in Midd1,7- ton, U9NN J.J, February 1977: s;:rin.= on Ped Hill. USNM 647279, J. 5. Lance. _ 7 . August 1977: EP '-' i7= on V4k7,=tsida o-F Middlton, HENi1 647244, J. 5. Landye, 21 February 1977: Sam Adair Sprinc, US NM 84724f. J.j. Landye. 21 February 1977: ANSP 12112 (holot!- e. API=ON. - MOHAVE COUNTh•: iN=t E n4; US-- N;.f E4720. 5.7. L-7Andys and G. Etards. 27 February 1777; si=ring uPstream Virgin River -F rom tt1 aid. i!9NN 9472.:!2. J.I. August 1977; NW of Litticm-fi i. W. Rinne, 17 November 1975'0.

DIAGNOSIS. A small sPecies having a globose to ovate-conic

hell. Penis with enlarged filament bearing (on its dorsal surface) elongatr= glandular ridge; ventral genial surface with two to three additional ridges. Holotype (Fig. 10a) height, 2.16 mm; width, 1.57 mm; whorls, 4.0.

DESCRIPTION. Shell mm high, with 3.5-4.25 whorls. varying in shape from globose, with well-rounded. unshouldered whorls and relatively large (60% cf shell height) aperture (Fig.

10c,d) to ovate-conic, with fairly flattened whorls and smaller aperture (Fig. 10a, b). Shape variation evident not only among, but also within populations, with enlarged conical specimPms sometimes forming a distinct size class. Sexual dimorphism significant in one of two populations studied. Aperture often partly or completely loosened from body whorl. Umbilicus either ab sent or chink-like to entirely open.

, Mr--l nir pigmentation of head-foot variable, dark throughout

(except for tentacle and snout tips. and sole of foot) in some populations, but consisting of light dusting on all surfaces or even absent in others. FPnix..1 filament Pigmentation ranging from dark throughout to re r-An nt.

Ct,=ni7ii:4.1 filaments few. 7-10. Radula (Fig. 11 characterizeri by rslAtively few CUSPS on central and marginal t==th. r.=1,--iti4=.1v =mall (31% of body length); prostate relative large

( -70% n-F body length). Fenis Elongate (Fig. 13), filament 74% of penis lengtn. extending b=ynnni tip nf oenial lobe. anr- thickened (lengthiwidth. 42%). Lobe relatively sm.=.11, sometimes tapering. Glandular roes. Ridgs on ventral =urf.-w= positioned on or just behind penis? lobe. Bursa (Fig. 9,-) enlarged (84% of albumen gland length). F,=min,=,7 recetacle small

(407.) relative to bursa length.

REMARKS. This 1.47Lriles, does not closely resemble any other

Arizona but is similar to an undescribed sgezies rom A Virgin River drainage, southwestern Utah.

TYPE LOCALITY. Pilsbry (1916, P. 111) gave the type locality as "Washington County, Utah." We suspect that the spring in

Middleton (USNM 847210) may actually be the type locality as specimens from the spring very closely resemble the holotvpe.

DISTRIBUTION. Springs along the Virgin Fiver in southwestern

Utah and northwestern Arizona. PyrguloPsis b=tcchus Hershier. new sPecies

Grand Wash sPringsnail

Figs. 9d, 10e-g, lid-f, 12b, 14

1 FT.7 .77-2110E-11 Willi=ims et al.. 1=Sf: 72.

MTERIAL EXAMINED. ARI 7ONA. MOHAVE COUNTY: Grapevine Spring. URNM 859077 (hmlotygP), s-io78 (4 p=.r:ItypPE), S47203. J.J. Landye. 4 June 198Ci.

DIAGNOSIS. A largP.-=i7PH. o• - e1 led sPecies. FPni= =mal1. with thickened. unPigmented filament; Penial lobe enlarged and oftPn bifurcate. Cne to three glandul...r ridges on ventral Pc=ni.A1 surface, a1on die.tal dgP of penial lobe. Hclotype ;Fig. .1.De, height. 2.41 mm; width, 1.78 mm; whorls. 4.0.

DESCRIPTION. Shell (Figs. 10d-e. 12b; height, 2.3-3.1 mm. with

4.0-4.25 moderately rounded. un=hmuldered whorls. Sexual dimorphism not significant. Aperture usually separate from body whorl. Ilmbilicu.=

Head--Foot usual iv dark iv pigmented throughout, exr- ept ior tentacles and a broad central patch on sides of head-foot, which are either- lightly dusted or unpiqmnted.

CtPnidium relatively small (19% of body length). with 16-71 filaments, Radul.=. (Fig. 11d-f) with relatively few CU=4DS on lateral teeth and numerous cusps on cuter marginals. Testie large, filling 54% of body length. Penis (Fig. 14) stout.

Pen id filament extending bPvond ti= n-F penial lobe. width cf filament/length, Glandular ridgee, 1-7. Capsule gland

( F i g. 9d) small (76%) relative albumen gland. 9.Pmin=1 receptacle only 42%. cf bursa length. This species has a distinctive penis =hah.= and glandular ridge pattern that is most similar to those of F. conicus, 444404.441WeAt.

TYPi= LnCALITY. Grapevine SPrinc3, Mohave County, Arizona.

DF:-TRIEUTIPN. Gr=tPPyin= and Whisky Springs in r;r=nri Wash, north of Lake Mead.

- ETH ICLOGI. Nam=d aft=r E•acchus. th= 1: r: n . (V,fis avizoAriCai reforrimg

-, the Grand Wash E.7 rings,

PyrguicPsis cdhicus Harshler, new species

Kingman springsnail

FiE. 1 t-. 1=, 17

Trvihni= n. SP. Landye, 177:: 42

MATE-UAL E

DIAGNOSIS. A mcderate-sized species, having an ovate-conic shell. Penis with short filament and single glandular ridge near tip of penial lobe. Holotype (Fig. 15a) height, 2.67 mm; width,

1.69 mm: whorls, 4.25.

DESCRIPTION. Shells (Fig. 15a-d) varying from 1.8-2.7 mm in height. Whorls, 3.5-4.5. either well rounded with distinct shoulder or only mildly convex, without shoulder. Sexual dimorphism significant in one of two poPulations studied. APERTURE OFTEN LOOSENED FROM BODY WHORL. Umbilicus open.

HEAD-FOOT usually without pigment, or with light dusting on

, =FLOUT; c- uhepithelial pigment absent. Penial filament

pigmentation light.

Ctenidial filaments, 17-21. Central radulAr teeth (Fiq. 16) with only four anterior rusp,5= en either side of enlarged. central

CUSP. Penis (..Fig. 17) elongate: filament half as long as Penis, sometimes extend in bevond penial lobs. Fenial lobe simpl= and

with =ingl.= glandular ridge positioned at Or just

proximal to tip of lobe on ventral surface. Less than half (42%) of bursa posterior to albumen gland (Fig. 9e): seminal receptacle

53% of bursa LE=NGTN.

REMARKS. The somewhat elongat.= shall of. this =Pi='""=.= 1=H to

its initial assignment to Tryonixt by Landv (1977) .

TYPE LOCALITY. Dripping Springs, Mohave County, ;:l riodna.

DISTRIBUTION. Dripping, Cool, and Burnsil rings ro==r i. in.7,man. p ETYMOLOGY. From Greek konikos, referring to the conical shape of the shell in this species.

24

•

- • , Pyrgulcgsis morrisoni Hershler, new species

Page springsnail

Fig!. 9f, 15e-h, 1E, 15

Fontalicella sPF. Williame, 15E5: 15.

MATERIAL EXAMINED. ARIZONA. YAVAFAI COUNTf: Ta/asci 3=rings, UE.NM 84=0, 5.J. l and%m and C. Emlk. 2 nr.to-ler 1977; Face Scrings, LIE,M Pctr=t-rme-) 4 , L G 7271, J.j. Landc. e and • Eml. Onto ‘mr

DIAGNOSIS. A mecium-sizmd =17,c,ri==- with ovate nr ovate-conic mhmll. Penis with narrnw filAment of medium length. penial lobe relatively 14'ge. Sin le =14ndular ridge Pcsitionca on ventral surface near ti P of pmni41 lote HclotyPe he

2.00 mm: width. 1.45 mm; whnr1'4, 4.0

DESCRIPTION. Shell (Fig. 15e-h) 1.E-2.5 mm tall. with 7.75-

4.50 slightly convex whorls. Sexual dimorphism significant in one of two populations studied. Inner lip thin and usually

4 pr 7=md against bodv whorl. Aperture less than half of body whorl height. Umbilicus oPmn.

- Pigment either absent from head foot, or consist np of light to moderate dusting throughout (except tentacles). Fa,nial filament either pigmented along entire length (sometimes darkly) or without pigment.

Ctenidial filaments, 15-15. F.,idula (Fig. 1S) distinguished by large numbers of cusps on central and inner m4rginal teetn.

- Testis filling 46 54% of body length. Penis (Fi. 15) elongate; lobe both wide and long; filament only rarely extending beyond genial lobe. All specimens seen had sole glandular on ventral surface lobe. Seminal receptacle EG-105% 01: bursa lengith.

FE'IARKS. The distinctive Peni= of this sPecies. with a 1arg7e lob*. slender filament. and single glandular rice, is most

Similar' to th.It of F. azhcariceerei..-

TYFi= LOCALITY. Fage Eprings. YavaPai County. Ari=cna.

rII7TS12UTInN. arc several ' the. 5 in northern Verde.

ETYNOLOG'r.. Named aft=r the late Dr. J.FE. Mcr!--ison his czntributions to tha etudy_ m_ f Hyrirobiir,a=.

re,A E7E.:ES

Montezuma Well s„-ringsnail

Figs. 7=, d. 21,

7 Fcntel joel 1-a s. Williams et al., 15:55: 1 ..

MATERIAL EX AMINED. ARIZONA. YAVAFAI COUNTY: Montezuma Well (swallet). USNM 55904: (holotyge), S59044 (3 parstymes). 547233. J. J. Landye and G. Edwards, 10 May 1970: Montezuma Wall icanal at outflow), USNM 547270, J.J. Landye, 10 May 1970.

DIAGNOSIS. Shell ovate to ovate-conic, medium-sized, with slightly loosened whorls. Distinctive dark pigmentation on snout. Penial filament short, lobe enlarged. Dorsal genial surface with two elongate glandular ridges (one on base of filament), one-two additional ridges on tip of lobe and on ventral surface. Holotype (Fig. 20a) height, 2.65 mm: width,

1.75 mm; whorls, 4.5.

DESCRIPTION. Shell (Figs. l2a, 20a-c) height, 1.7-2.7 mm:

C whorls. 3. 5-4 . • highly convex with indented sutures. Loosening

26 of whorls along axis noticeable in most specimens. Body whorl only 77-74% of shell h,--- ight. Sexual dimorphism significant in one of two populations studied. Aperture relatively wide. Inner lip thin, and either just touching or with much of lent h +used to body whorl. Umbilicus open. UPerculum (Fig. 12d) ovate.

Entire snout, distal half of snout. or just oral leoPpets with very dark, purple pent. Ti=nt.=c1==. sides of h=ad/foot, anterior . edge of foot with =mm=wh.=t lig,ht=r pigment. Penial filament unpiPmented or with small pipm=nt patch near bas=.

Ctenidial filaments. 15-19. Lateral ans!les on central tooth widely diverqing (Fi7. 21). Central and marginal teeth with large numbers of CUSPS. Glandular ridges, 2-4. P=r1-;.= moder=ttely elongate. Filament only 70% of peni,=e1 lsrqtb. rarel? extending beyond penial lobe. Width of filament 407. of length.

Two elongate ridge=e on dorsal surface positioned on or just proximal to filament, and near center of penis- In addition to elongate ridge along (ventral) edpe of lobe. =mA.11 ridge sometimes present on central portion of ventral surface. Seminal receptacle 647. of bursa length (Fig. 9p).

TYPE LOCALITY. Montezuma Well, YavaPai County, Arizona.

DISTRIBUTION. Endemic to Montezuma Well and its upper outflow.

ETYMOLOGY. Referring to the type locality. PvrguloPsis solus Horshler, new sPecies

Brown springsnail

Fig. 70n-f, 23, 24, 25a

MATERIAL EXAMINED. ARIZONA. YAVAPAI COUNTY: Brown Spring. USNM E59045 (holotYpe), 359046 (3 paratyPe=.), 850290, j. j. Landye, 4 February 1936.

DIAC;NnIS. Shell ovate. small. Penis small. with , r11,:krgr..,d,

bifurcate lobe. Elongai!e glandular ridge along lobF'= edg,

ridge on ventral surface. Holotvpe (Fig. 20d) height,

1.69 mm: width, 1.29 mm: whorl-7, 7.5.

DESCRIPTION. Shell height, 1.4-2 mml whor1.7, 3.5-4.0,

moderately convex. Body. whorl relatively large (7.i77% c =h47:11

height). Sexual dimorphism significant. Inner liP moderately

thickenPd, fused to or slightly separated from body whorl.

Umbilicus open.

Pigment on head/ foot may be absent, or range frcm very light to

quite dark dusting on all cur-faces. Fenial filament darkly

pigmented.

Otenidial_filaments few, 11-14. Radula shown in Fig. 27.

Penis (Fig. 24) almost square in shape and very small, not

extending beyond mantle collar. Filament positioned at only two-

thirds of penis length, moderate in 1. .higth (46% of penis length),

and rarely extending beyond penial lobe. Width

filament/len!=ith, 40%. Bursa (Fig. 25s) onlv 76% of albumen gland

Seminal receptacle 83% of bursa length.

RE1ARKc2:. No close reltionshiP is apparent between F. solus

and other Arizona congeners as the minute, squat penis of thi=, sPecies is unigue among southwestern FvrquloPsis.

28 TYFF LOCALITY. Brown Spring. YEV2Pai County. Arizona.

DISTRIBUTION. Endemic to type locality.

ETMOLUGY. From L,Rtin meaning alcnC or single. and

rFfet-ring to the endemi=.m of th1=, =r'i== in a single sgring.

FYrgu1t777Ei.= =immi.=x HP.r=hi=r. neN amdciaa

Fdsail sprinanail

Fima. J'3 3.

FORTAIIRELIA =p. Williams et al., 19E5: 19.

7 MATERIAL EXAMINED. ARI ONA. YAVAFAI COUNTY: Edsail Smrim57s. UENN 847222, J.J. Landye and D. &elk, f "'l ay 1972. GILA COCNTY: , =mrinc; near Etrawb=rry. HENM E59C.)49 (hcidtyme'). 8:90f0 (4 Paratypes'i, 847=6, J.J. Landy,=. lA Cictdder 1971.

DIAGNOSIS. H moderate-sized F-mEC1=7 with ovate-conic =nr=ii. Penis with narrow filament of moderate length; single glandular ridge on ventral surface of genial lobe. Holotype (Fig. 20g)

height, 2.41 mm; width, 1.47 mm; whorls. 4.5.

DESCRIPTION. Shell height, 2. 0-2.5 mm; whorls, 3. 5-4. 25. unshouldered and moderately convex. Shell relatively elongate for genus (shell height/shell width, 150%). with some sPecimens conical in shape. Sexual dimorphism significant in one of two populations studied. Aperture usually separate from body whorl.

Umbilicus open.

Snout ungigmented, sides of head/foot with light dusting of melanin. Penial filament darkly pigmi=nt,=d along virtual entirety of length.

Ctenidium relatively large (27-31% of body length), with 16-1g

29 filaments. Radul; =hown in Fig. Penis (Fig. very elongate. Filament 57% of penis length. Projecting well beycnd tip of penial lobe. Width of filament only 2E of length.

Penial lobe simple; single glandular ride positioned near di=tal edge of lobe, variable in size. Ovary only 12% of body length.

Over half (5%) of bursa posterior to albumen Seminal receptacle 6,5% of bursa length.

FENARKS. P. =inPl=x m.A.iv be clo==lv allied with other Arizona

- - having a relatively simPle penis. including P. conicus.

P. mnrri=r,.ni. and F. =anc=riosensis.

TYFE LOCALITY. Spring near Strawterr' Gila County. Arizona.

DISTRIBUTION. Restricts to s number of springs feeding Foesil

ETYMOLOGY. From Latin =impl=x, referring to the simPie condition of the penis, with few glandular ridges.

30 Pyrgulom=is cmnilitentis Hershler, new sPecies

Three Forks springsnail

Figs. 2d-q, 25c, 2E, 29

N. gen., n. SP.. Landye, 1973: 47.

N. pen., n. sp. Landye. 1981: 27.

- MATERIAL FXANIN): . ARI2ONA. AFArHE COUNTY; sPring on N =ide of Blanket Creek at Throe Forks, U9NN ? 57053 Cholotv=s). EF90F4 4 ParatyPesJ, 847274, J,J. L=tndvP. 15 September 1977; =Pring at Three Forks. USNN 847279, J. J. L=ndYP, 7 October 1973.

DIAGNOSIS. Email low ovate:, to ovate-conic. —variable in

Penis with small filament. elorm.f= lobe. and oni=rg=t ventral accessory crest. Glandular ridges numerous on Ohth peni=1 surfaces. HolotyPe (Fig. 2d) height. 2.E0 mm; wimth.

1.9/, mm; whorls. 4.5.

DESCRIPTION. Note the sh=11 size and shape differencz amonc, the two known populations -For the species (Fim. 2O-g). Shell height, 1.5-4.E mm; whorls, 3.5-4.75, unshould=rm and well- rottndod. Sexual dimorphism significant in one of t o populations studied. Dark, thickened periostracum covering entire =h=11 surface. Inner lip thin. limbilicu= chink-like to open.

Dark pigment on snout and tentacles characteristic, pigment somewhat lighter on =iri,==. 0-F heat/ oot. Melanin pigment sometimes also scattered over much of penis. Subepithelial pigment dark in proximal half of p=nial filament, sometimes extending into penis as well.

Ctanidial filaments, 17-23. Ctenidium only 19-21% of body length. Radula shown in Fig. 26. Inner marglnals with as many or more CU5P5 as outers. Testis 46% of body length. Penis

31 •=iongate (Fig. 29); filA.memt only T. of psnis length, not projecting beyound tip of penial lobe. Filament relatively stout at base (width/length, 47%). Lobe sometimes with narrow, terminal process. Accessory crest on ventral surface swnllen.

Glandular ridges. 4-6, rarely enlarged or elongate. Otie ridge

(sometimes absent on proximal (dorsal; half of filament: t c on dcr l portion of lobe (or just proximal tc lobe); one just proximal to tiP and one-two on crest. Ovary only ll% o+ body length. Oviduct C",cil COMi LSX. coil diamoter only 30% of albumen gland length (Fig. Hal -f of bursa Posterior to albumen gland. Seminal recePtacl,.= 7)% of bursa length.

REMARKS. This species link= with the P. glanduicsus-group on the basis of penial morphology.

TYPE LOCALITY. Spring on N side cf Blanket Creek at Three

Forks, Apache County. Arizona.

DISTRIBUTION. Several springs feeding headwaters of Black

River.

ETYMOLOGY. From Latin confluens, meaning a place where streams meet, and referring to occurrence of this species at ThrP=,

Forks.

32 PyrqL.,inn=i= PancP.rioPensis Herahler, new spacies

Evlas sPringsnail

Fig.. 1 7e, 25e, 70a-gi, 31. 72 iii.,44teassi4.%ydroA i FON7`alieell4 ZS,?. eFFe guAl 1454, /4T3: 36T aP. J± 11 ms Pt al., 1 E: 14 4 il ro..44/d444/14. /60 46444 44LN ELVTD.,41 /4ETC; /EQ.

MATEFJAL EXANINED. ARI2ONA. 6RAHf:1i FOUNT i =primq neP.r 2v1sa, UENN 247217. J. 1. Lardve, 20 Nay 1'5717 spring near Gila RivPr bridq NW flf UENM J. J. Landys, 1'371; large sgring N of Ev1.4.=, 11"-7NN J.J. Lamdve srd D.W. T=kylor, 20 APrii 1371: springs W Tom Niece Springa. UENM -7-.790f1 (holotyPe, E7902(4 , 84722, J.J. Land _. J!inP 1'577; sc==== W c4: Tflm '5.prins=, UENN 5:4722f, J.j. L=ndvP, 27 June 1577.

- Dif-;GNCEIS. Shell varying in Phpgf= frf!rn gITJtose tc

elongate-comic. Feria masaive, with eionsate filamemt and thr==

slandular ridsea at most. seminal reda acle atPPnt.

nE=j7RIPTION. Dverst in shell size and shape shown im Fig.

, 30a-q. Enth shell -forms sometimes present IN single population.

with conical specimens usually enlarged (often female). Shell

height, 1.1-2.4 mill; whorls, 3.25-4.25, moderately rounded.

Sexual dimorphism not significant. Small globose specimens with

relatively large apertures (50-55% of =hell height); larger,

conical shells with smaller apertures (48% c-F, shell height).

Aperture usually separated, sometimes greatly so, from body

whorl. Umbilicus open. Operculum shown in Fig. 12e.

Snout darkly (black) pigmented, tentacles and sides of

head/foot with lighter piqment, central section of latter

sometimes unpigm.=nt-,==d. Fenial filament either darkly piqmented

along entire length or unpigmented.

Ctenidial -Filaments, 12-20. Radula shown in Fig. 31. Penial

filament relatively narrow (width/length, 26-25%) and elongate

33

(67-84% of penis length), but usually projecting perpendicular to penis length and therefore not ext,=nding beyond tip of penial lobe. Enlarged lobe usually untapered. Glandular ridges, 1-3.

Dorsal surface of penis with or without single ridge; ventral

=itrface with single enlarged ridge and sometimes an additional smaller one. Capsule gland (Fig. 27:7. ? consistently lonqer than albumen gland. Oviduct coil di A met Er 1;Irge (G1-22%) RE.STIVE TO

- , albumen gland length. Bursa 60-3S% of albumen gland 1 =NGTH.

Note th-rtt duct EXITS LATERAL TO bursa tip.

-, REMARKS. The loss of seminal r=r =PTACIE in this species is not known for any 41110 congener.

. TYPE LOCALITY. Springs W of To Niece Ea-rings, GRAHAM COUNTY,

Arizona.

DISTRIBUTION. SPrings along Gila River between Bylas and Pima. 10444 ETYMOLOGY. Named after the San Carlos Indian Reservation near A E 1E-E. Fyrgulmpsi,=. cmchisi Her-shier. new sqecies

San BernA.rdinm sgring=41,,kil

Figs. 25d, 30h-k, 33, 34

FONTLICELLA sP. illiams et al., 17, : 50. MEL': sPr.cx,444/4“aa" 16,44164,4442" E. DE 14 F. MATERIAL EXAMINED. ARIZONA. COI-HIT,E COUNTY: Sgrings at an Eprnardino Ranch. Us7NN 8SC..5 (h,71!--.tvPi=). Ef.7056 (2 p.=tratypes!..

DIGNOSIE. A small-sized species ha,iing ovate-conic shell.

Penis without lobe; -fil...tm nt Dorsal and ventr51

..;,. wfaces each with single Plandular rids. HolotyPe

heigint. 1.af mm: width, 1.04 mm: whorls. 7.5.

DESCRIFTION. Shell ho=ipht, mm; whorls,

sI ightly rhnvg.x,, with sutures flmly slightly intenteo.

Sexual dimorphism siF„nificant. Inner lip thin. Umbilicus ogen.

Snout with light to dark dusting of pigment. Sides ci

head/foot somewhat less pigmented than snout. although

operculigerous lobe often quite dark. PENIA.1 filament darkly

pigmented along entire length.

Ctenidial filaments, 12-15. Anterior portion of cntral tooth

with relatively few cusps (4-1-4) ( FIG. ? ?' Inner and cuter

marsinals with roughly same number of cusps. Prostate only 11%

of body length. Penis fairly elongate (Fig. 34). Penial

filament 61% of penis length, extending distal to anterior tip of

penis. Filament three times as long as wide. Glandular ridges,

2. Ridges positioned just proximal to base of filament. Capsule

gland slightly longer (108%) than albumen gland. Seminal

receptacle body minute, length of receptacle (body plus duct) 67%

35 of bursa lengtn.

REMARKS. P. ccchisi is most similar to P. thomPsoni. which also have small Penes with reduced penal lobe (absent in the former) and single glandular ridge.

TYPE Lni71=1LITY. Spring at San Bernardino Ranch. Cachise County.

Arizona.

DI3TRISUTION. 7%nd othf=r

at Ermi Bern,z.rdinc

ETYMOLOGY. Named a-r:t2r Cochise Ccunt.

tHicmpsr,mi Her=his,-, n51-4 EPECISS

I-Har- hur- =t Eprinqsmail

Fiq=. , . C2Ndz4e44460e.../(yd1 ,4; ;a(.." /44444444 /9:69 6. FOntSliCella SP. Beguaert and Mil ier 1977; 214.

Fontelicella spP. Williams et al.. 1 S5: 19. 7444-IAI4IA•‘ ILT41444444-4 EA4D Al MATERIAL EXAMINED. ARIZONA. SANTA CRUZ COUNTY: Cottonwood SPrings. USNM 647221. J.J. Lanove and D. Belk. 1 7 August 1977; Monkey Spring, USNM 647223, J.J. Landye and D. Belk, 2 November 1969; Canelo Hills Cienega, USNM 647226, J.J. Landye, 14 November 1976; Sheehy Spring, USNM 647235, J.J. Landye, 16 April 1976, USNM 647227, 14 November 1976; Peterson Ranch Springs, USNM 655057 (holotypPO, 653056 (7 paratyPes), SUNORA (MEXICO). Ojo Caliente, USNM 847251, J.J. Landye and D.W. Taylor, 7 May 1971.

DIAGNOSIS. A moderate- to large-sized species having an ovate or ovate-conic shell. Penis with elongate filament and single, small glandular ridge on ventral surface of penial lobe.

Holotype (Fig. 35a) height, 2.57 mm; width, 1.67 mm; whorls,

4.25.

DESCRIPTION. Shell (Fig. 35) height, 1.7-3.2 mm; whorls, 3.25-

36 M m(nderately convex with slight shoulder. Sexual dimorphishl significant in two of four populations studied, in nni= of which males are larger than females. Inner liP thin. Aperture fu=..---=.d to or separate from body whorl. Umbilicus chink-like or open.

out and anterior portion in+ foot (dorsal surface) with light to faIr li dark pigment. Remainder of head/foot usually i mpigmented. Penal filament darkly Pigmented throughout.

filament=.. 11-20. Radula shown in Fig. 36. Testis

length. Frostmt.= only 7-q% of body lenth.

Penis =.guat to elonsate. Penis filament centered at RO-97% of penis length. Filament 35-103% of Peni= D=ngth. and only slightly thickened (wirithilPngtn. 27-4). Penis lobe simple.

Glandular ridge usually located nixzer ti m of lobe. Oviduct coil relatively complex in that initial turn of iCOP bends back on itself (Fig. 25f). Most of bursa length (55-85%) Posterior to albumen gland. Seminal receptacle 53-72% of bursa length.

TYPE LOCALITY. Peterson Ranch Spring=, Santa Cruz County,

Arizona.

DISTRIBUTION. Half a dozen springs in Southern Santa Cruz

County. Arizona, USA, and northern Sonora, Mexico.

ETYMOLOGY. Named after Dr. F.6. Thompson, in recognition of his contributions to study of North American Hydrobiidae.

Genus. Trvonia Etimpson

REMARKE. Hershler and ThnmP=on (in press) provided a detz.i 1a

description for the genus. Hvaloayrque Thompson was considri=d

synonym TrvoniA.. Trvoni.A have an Plonsate-conic to turreted

pg-ni=- with a small numoer of plandular per al

ovoviviParous mod,=, reProduction, recePtidn in the

-F emale via a short spi=rmth.,,--nal -imr- t. The genus has a dis:';unct

codurrinp in much o+ Florida as well a larse

Fart the arid Eouthw5=t. These snails are relat;thd to a sroup

oi genera having elongate-conic shells and mammi.form

, 7 - slants. The genus nnnt=tins a numbFr 7" ilndg-soribet E7SC1='-: -F rnm

the Southwes7; as w..--11 as ten descriteO spedies.

Trvonia pilae Hershier. new sec lee * Gila tryoni;i

Figs. 12c. f, 38a-e, 39, 40a-c, 41, 42d-i, 43, 44, 45a. 46b

14 14 7yo41er 444 ý ,1/A-ta /FT3 '369 A1I.4.464 E44 EDS41/4/ Trvonia SP. Williams et al., 1985: 15.

AI MATERIAL EXAMINED. ARIZONA. GRAHAM COUNTY: small spring near Bylas. USNM 859055 (holotype), 855060 (4 paratypes), 647257. J.J. Landye, 30 January 1971: spring near Gila River bridge NW of Bylas, UENN 847255, J.J. Landye, 30 January 1971; large spring N of Bylas, USNM 847258, J.J. Landye and D.W. Taylor, 20 May 1571; springs W of Tom Niece Springs, UENN 647254. J.J. Landye, 27 June 1573; seeps W of Tom Niece Springs, UENM 647252, J.J. Landye, 27 January 1973; Cold Springs, UENM 647253, J.J. Landye, 27 January 1573.

DIAGNOSIS. A moderately large-sired species having an

elongate-conic to turreted shell. Cephalic tentacles with well-

defined linear tracts of hypertrophied cilia. Penis with two

glandular lbes on inner curvature near ti P and enlarged lobe on

outer curvature at base. Outer curvature distal to lobes with

38 hl,pertrophied ciliary tuft. Bur=;:t r=i=ttively large. Hclo.yPe

(Fig. 3Ea) height, 3.25 mm: width, 1.4:'17, mm; whorl=, 6.0.

DESCRIPTION. Ehell (Fig. 3Ea-e) c1eai transg=tr=mt, and

5 without periostrarf!m. - hell often twice as tall a= wide, heqh t.

1.5-3.3 mm; whorls, 4.5-6.0, slightly to moderately convex with

indented sutures. Sexual dimorphism pronounced, with

larger than mal==. Prot000ncn (Fig. 12c =month and flat

;sometimes slightly degre=sed). with 1.0-1.25 whori=. Body whorl

57-74% of shell height. Aperture SiMPle, occupying less than

half of shell height, longer than wide. slightly angled Allov=, and

. rounded below, and either fused to Cr segarate from body wnrrl.

Inner lip fairly straight. slightly thickened and refiectet:

outer lip rounded and thin. Apertural plane slightly tilted rrm

coiling axis (adapically somewhat advanced) with long axis

inclined 25-30° to coiling axis. Umbilicus open. Telerconch

with strong growth lines. Operculum (Fig. 12f) amber.

paucispiral, over one and a half times longer than wide, and with

three whorls.

Snout (Fig. 39) longer than wide, terminating with -Fleeny oral

, lapP =te. rePhalir tentacles (Figs. 3. 40=..k) rirrow, slightly

expanded at tip=. and moderately elongate (only slightly

extending beyond snout ti7). Ciliary tracts on tentacles shown

in Fig. 40a-c. Snout and "neck" with scattered ciliary tufts as

well. Elac esPots located in slightly bulging_ cuter edges o-F

tentacle bases. Foot elongate, with anterior edge almost

straight and posterior portion expanded and rounded. Pedal

glrld=, 12-1S. Head/foot lightly dusted with epithelial melanin throughout.

- except for tentA.. 1 =. Narrow strip extending down of head/foot very lightly pigmented or unpigmented. Dorsal surfaces

of style sac and stomach darkly pigmented. Ovary with light

dusting, testis very darkly pigmented. Section of pallial cavity

, roof aoc'.ie the ct,=nidilLm drkly immg.nt.=H in males.

Penis with dark i mCr a ch naar ti see Hershier 1 :

44) =tr k.= lodes

and f=1 ,,-=wher= or dorsal surface. subopithelial pigment

chnc=mtr d on right a e mntle collar. Large. dull

- , - - glandular clusters in un!... k" 7- roxim.-t1 to tentacles. i2a.E-Ea" 01

tanaciea with white glandular clusters.

Ctenidium ;Fig. 41. f7t) eionciata. filling most P AL LIAI

cvt lenth. Fiacnant Uf ...r, atUv

Osphradium (Fig. 41. 0=-) =mall r=.1ative to ctenidium length. • Kidney (Fig. 47a, Ki) with small opening fringed by distinctive

lips.

Buccal mass small relative to snout. Radula (Fig. 42d-i)

typically taeniosiossate, with broadly trapezoidal central teeth

having two pairs of basal cusps. Lateral angles of csntrals

widely divergent, narrow basal process characteristic. Central

teeth with enlarged central cusp; central cusp of laterals

sometimes not enlarged (Fig. 42q). Inner and outer marginals

similarly shaped; outers with only a few more cusps than inners.

Stomach (Fig. 4:b. Ast, Pst) and style sac (Sts) about egual in

lensth; the former having a broad caecal chamber (Cc) along its

posterior margin. Posterior stomach chamber with single opening

to digestive gland (Dgo). Digestive gland covering posterior

40 half. of stomach and Pxt-,..- nding (ventral to testis) almost to tip of vi=ceral coil.

Testis large, overlapping most of posterior stomach chamber.

Frostate (Fig. 43a, Fr) small, flat, near-circular ir outlinf=, positioned 1a el or totally posterior to Pallial cavity. n,-=..f,=rns enters posterinr nnd of pro=t-4.te. and exits near anterior tip.

Fenis (Fig. 39, Fn) flattered, ;=longat.=. to snout, extending forward +rnm Att.-,:nhmnt witnnut coil ns.

Two smal I penal istes (F1, Fig. 45A) on inne curvature centereC

about two-thirds of ST outr curv.--rt tIkr massive. and usually fnldir-d .=t ginst IS near SSC. il

lobes bearing mammiform glands Tip cf with Pronounced swell inc on inner curvature (ics. Fig. 45a. Vas deferens undulating in penis. terminA.ting in eversible . ilia

lateral to swelling. Penis surface, especially at ba-,4.=, well endowed with dense glandular clusters of several types. Penial ciliation shown in Fig. 44. Dense tuft of cilia at penis tip

'extending along outer curvature to level of base of distal lobe. and along inner curvature to distal end of swelling (Fig. 44a,b).

Proximal to enlarged lobe, three-four distinctive strips of cilia cross penis surface (Fig. 44c,d). Remainder of penis surface with scattered, small ciliary tufts.

Ovary very small relatively to body length, non-lobular.

Capsule gland (Fig. 39a, Cg) thin-walled, enlarged (covering entirety of style sac), serving as brood pouch containing three-

four shelled embryos. Embryonic shell (Fig. 39a, Em) with as

41 many as 1.5 wnorls. animal dark y pigmented. Capsule gland with narrow muscular sPhinct=r (Fig. 41, Cg) at anterior end; and posterior bend (Fig. 4eb, Cg) which extends forward (partly ventral to distal section of caPsule gland) almost to end of pallial cavitv. Albumen gland (Fig. 46b, Ag) comprising very

=mall. narrow. section of P;A1ii,R1 ovi.uct. disaPpearinc3 vntr.,11

, to bursa cosulatrix ( Ei0 to r,=,f-7-iv .--vidur- t. Oviduct (f-i\, with

, '.. impie coil ( ventr.ai to bitr=.= pr,-ximl to r.1,,Pnin9 to altumcm

. ,- ...qiir ... loi 0.72 Iv). Bur= rlub-=haPed, s;nall relative to p•=t1li=t1

-. rvid!u t, and larqely otsc!!red anteriorly by albumon 91and. 2ursa gradually narrows anteriorly to giv= rise to =Perm=17heri ONct

C77 just posterior to end o4: 9allial ora y. Eenin,rli rebec=tacie

C-=,r) minute. with short duct cr--nnr- tin= to oviduct. (i-ftEr this

juncture, duct (Liar.' coils several tines ventral to bursa befhr=

looping back to open into dorsal surface of bursa at or just

proximal to origin cf spermathecal duct (Osr). Spi=rmathecal duct pressed to ventral surface (colitmellar side) of capsule gland.

Duct narrow, non-muscular and short, opening just anterior to end of pallial cavity (Fig. 41).

REMARKS. This species is not very similar to either T. guitobaguitae from Arizona or undescribed taxa from New Mexico

4 (Taylor, 1983).4114:1ts affinities may lie instead with Nevada species having relatively many penial lobes, such as T. clathrata

(Hershler and Thompson, 1986).

TYFE LOCALITY. Small spring near, Bylas, Graham County, Arizona.

DISTRIBUTION. Springs along Gila River between Bylas and Pima.

ETYMOLOGY. Referring to distribution of this species along

Gila River.

42 Trvonia guitobaguitae Her=hier, new smedios

Quitobaguito tryonia

Figs. 3Ef-h, 40d-i. 42a-z. 45a, 46a

Trvomia imitator (Pilsbrv) Esclua,=rt and Miller 1977: 213.

MATERIAL EXAMINED. ARIZONA. PIMA COUNTY: Quitobaguito Springs. USNN etTc-:o61 (holotype), (4 ParatyPes). 647256, J.J. Landye. 13 March 1971.

DIAGNUSIS. A minute-sized specISS, with ovate-con to

,, turreted shell. CePhalic tPrit toir.,= without well-defined c iiirv

,. tisacts. Penis with two lobr-m=, on inner curvature. a sm.111 one just distal to midPoint and a =r;mgmwh.....t larqsr one near the base.

Much of distal portion cf penis with dense cover of cilia.

, Bursa very =mall rlativP to p;AlliA.1 oviduct. Holotymg: iFiP.

:36f) height. 1.96 mm: width. 0.9A mm: whorls. 4.5.

DESCRIPTION.. Shell (Pip. 3F-h) h,--- ipht. 1.4-1.P mm: whorls.

highly convex with deem sutures. Amertur-al liP thin.

Inner liP only = 1 P1--, t1,, refler- ted and usually fused to body whorl. Umbilicus chink-like.

Dark pipimentation on visceral coil near uniform. Snout and operculigerous lobe usually darkly pigmented, elsehwere headlioct having light dusting of melanin, with broad central section on sides of head/foot sometimes having distinctively light coating.

Penial tip with piqmont patch.

El-- ttered ciliary tufts on ci=phalic tentacles shown in Fip.

Central teeth of radula (Fig. 42a) with simple pair of

basal CUSPS. Lateral teeth (Fig. 42b) with enl.,-trged central

CUEP. Narginals shown in Fig. 421o 4 c.

43

penla snown in Fig. 4fb. Note thtvaa deferens undulate only slightly in penis. Extensive penial ciliation shown in Fig. 40f. Filiation extending on both cuter and inner curvatures to level of ba=,=, of diatal lob=. Eursa copulatrix comPlex shown jr Fig. 46a,b. Note the minute-sized bursa (Eu). which is not gr,=,.,tly ov,=,r1.4.pp.=.d by the ..=t 1bumpn

REMAP.W.E. The reduced lob:.ttinn fl-f. T. c.74 itoillauitas is shared by two undeecribed csn:7eners +.rom Utah Craylor.

- 1 FE LOCALITY. Quitotauito Sp'iiqs, Fla.O 0: Uflt. urizona. DISTRIPUTInN. iv Pe local I- , ant everai adjacent aprinqe. 2 - FTYNCi0F;Y. Rei:errinq to thP type C:

NORPHOMETRICE

discussed by Mayr (1=76;3) and (17.E1), .-,: =4,===m,-=nt 0+ systematic =.tstu,-7 oi ,z111 ,,p,LA trir without genetic or experimental data involves gauging population divergence using morphological criteria. Multivariate analysis of morphological data can contribute to the decision- -, making process in such situations (Wiley, 1981: 65-67 , ._)..J=..J= -0,,._,-, , and Pe therefore used 4g-04...etrT to examine patterns of variation among the 12 allopatric PYrgulopsis spp- described above.

We employed canonical discriminant function analysis, as this technique is particularly suitable when one seeks to evaluate distinctiveness of a priori designated species.

Unlike principal component analysis, discriminant analysis provides statistical evaluation of specimen classification.

Means for all raw counts, measurements, and ratios except one

44

(CSW, ff.mal,==) were hei;ercgeneoue among populations. Initial discriminant analyses indicated that significant ==pAration

(partial F-test, p<0.05) of closest groups (consisting of

topotypes of given species) occurred After stePwise selection of subsets of male and female anatomical variablese) ELection of additional variables no further separat4WlegrouPs and eventually (aftE r 1A-20 Added) result in decreased separation. To avoid possible confusion of pAttrn= enerated hv addition of 4.44=does 1.4044er variables, dierriminAnt amAlv =7 were eagia rerun using subsets G.11-.1,‘. = with t';TT-,... COnTiiStinq 0+

All variables selocted (in initial analyses) to tne point at dU4WAA.44, which consistent detween clasesr grouPs e. e.A.4444 The reduced variable ==I-= as

follows: male anatomy (11 variAblA,$), CTW, TS/EL, L=. GL, L7/L7,

LFI, WFI, CTL, LFR/PL. L3: female anatomy (13). CTL, E3.

01W, COP, CSW, FI, ASL, OV, ERL/BUL, GEL. BDW, OCT, DEL/CTL. For shell data (consisting of relatively f.,=,w variables), decreased

separation cf closest group'As occurr Ed aft E r every step in initial analv.-...ss and therefore tOree-e-runsefr-utili,-ing all variables-)ere

- used. Rbsults of discriminant analyse!) c on

.azz.t 4-6-er tre !mivcFp. in Tabls 3-, with scores on the first

two discriminant functions plotted in Figs. 47 and 42.

Correct classification to sPecies for grouped CZS2s.

, representing topotypes for each saciaa, was Ec7'% (overall) for

, the analysis based on male sh =11 d A ta (ranging from 80-100% among

groups), 93% for the analysis bA,=.=d on female shell data (63-

i :•;. and 100% for analy=s u=,-ing Anatomical data (Table 6).

45 The somewhat lower level of succeesful classification based on shell data is reflected in p lo te of scores on the -Fir7-.t two discriminant functions (Fig. 47)• which collectively

68.R% (male) and 78.7% (fi...male) of tntA.1 variance. Note that

. groups are not w=11c-eparated. esPeniA.Ily in the analysis of

1, , f=m..‘ = =h =11 data (Fig. 47b) in which ==,nh -.6.42-.-trw• is overiaPPed by

, at least one other. R.„grificant s=p,,r.=,timn of closest groum= (7.

F. simPlex: 3. F. ccrcus was not achieved ,=,v(=n Al.ftf=r all =tPp= wre comPleted .4...R-401+.11.-4~A-re4m1 (p=0.77): n:ra that all Pnint'.

- , rg-pre=enting the former =e1.24e.s are enclosed within the latterip

For the analysis nf male dat.=, (Fig. 47b , significant separation of closest groups was achieved (p=0.04), yet all

exhibit overia4except twci

Interpretation o+ the two discriminant functions (based r7n loadings and correlations in Table 3) suggests that function

, r=pr.===nt= sios in both (decreasing along axis in male

Plot, increasing in -f emale plot) whil= -f unction 2 correlates with relative hil heinht (increasing along axis in male plot, decreasing in female plot). Not= that most ....4-044.e.separation of groups occurs along the size-correla ted axis. The two non:- overlapped groups in the male plot (1. F. d==7.7,- rta, separated on axis II; 12, P. cochisi. saarated on axis I) are unusually globose-shelled and small, respectively. Classification of ungrcuPed cases was highly variable ' 7 - from 0% for several populations of F. thompsoni to 70% -for an additional population of thittspeciee.

46 suPerior clas=iiieatinn reeul ng from analyses based on anatomical data is reflected in relatively separated grouPs in plots of score :Podid: on the first two diecriminant functions (collectively exPlaining 64% Cmale3 and 67.6% tfemale]

total variance), with six (male) and three (female) grouPs

,-1 non-overl=pme Fi. 4Sa.b). Eignificant (p<.005) separ=tion closest groups VA S: SCMiEVeC after the final etepr -

In the Plot tesed on analysie male anatomical riate,

- =emaration of grnue= ie eeeri z*I'lrsci both axes

.- 1), largely interPre•tai.1e in term= of penial morpholcgy ant bocy•=iz=. Function he=

. , particularly high loadin-7,s of GLR and WFI (TA:L7, 1e 4). SPecies with low scores on this axis such as F. thomPecni (11) have PE711.1= with thickened filaments and few (usually one) glandular ridges

(see Fig. 37), while those with higher scores (9. P. confluentis:

8. P. glandulosus; 5, F. montezumensis) have penes with narrow

-filaments and more than two ridges (see Figs. 7, 8, 22, 29). The

axis has high loading... of other peni=1 variables (L2,

L3/-2,1 LFI) ae' well as = =ize cnmpcnent (note high loading of

EL). Species with low scores on this axis have relatively long

- 4 penial filaments and/or lobes (Figs. 13, C 7 ) (10, F. samcarloseneis; P. Eimple,-; 1, F. deserta; 12. F. ccchiE1), while species with high scores, such as F. bacchus (2), have shortened filaments and/or lobes 14). Note that ungrcuped specimene are fairly tightly clustered and positioned close to

toPotypes of their series (sepecialiv P. sancarlosensis). but

rarely overlap with them (Fig. 4Sa). For these unprouped

4.7

specimens, cl=.==ific=.tirin ranged from 07, ( two poPulaticns cf F. thompeoni) to 100% ( a third poPulaticm 04 the

In thethe plot based on an=lly=i= of fen=le data ( Fig. 4Eb group

=eparation is largely scores on th first

, discriminant function, which is clearl-y si==-cnrr=1=.t=.d. given high loadings of CTL, GEL, OCT, =.nd Si on it (Table 5). The sedond disdriminant function has high l oadin E. of OEL/CTL, EUW.

COP, and ERL/TF.UL. The partidiarly low scores of F.

- =anr- arlr==.m=i= ( 10) on this •=k x i= jJ result from absence of

CiL- SEMINAL recePtacie in th = sPedies. Ur:grouped specimens

( especially F. sancarlosensis and mcrris mi) ari= generally more seParated from appropriate t:-,p7tYp=-s than in tho= m=i=,

Also note the large variance .0-,f on tne first disdriminant fimrtion for sPecimens representing P. thompsomi. For unprouped r=t=e=. r1.===ifir=.tirin ranged from 0-4C%.

DISCUSSION

Results of the discriminant analyses support division of

Arizona Pvrqulopsis into 12 species, in that these appear well-differentiated, with discriminant equations based on four separate data sets for grouped topotypes nearly always correctly classifying

Discriminant analyses using anatomical data provide classifications Ltwat orz. slightly superior to those generated by shell data, and yield greater =.p=di=,= separation on plots

( especially based on male anatomical data) of the first two discriminant function scores, suggesting t4-yeed* the former data set

46 , ° discriminates these epecies somewhat better than THE latter ilioEmj

THIS MAY BE PARTLY DUE to differences in sample size: more shells

PER group were measured THAN animals (7-11 vs. 5), which 41.4.-404r

result= in increased variance in discriminant function scores -for' the former.

.7011-- SUPERIOR separation of groups based on 4.etti l. the first two discriminant furctione for male anatomical data r,.+1=-. -,- t=e0)1" relative diverettv ant of puantifioation of penial morpholo,Pv compared to =hell =.nd fem=de morPholosy. Morshologir- diversity amonc ehelle of congeners, ===.rt from size =nni pi=n ral form. is siisht (Fiss. •■•• 10, 1=, 20. 70. addition, =hell variables used. consietinp those

TR•=DITIONALLY EMPLOYED BY MALACOLOPISTS, are admittedly unsuited

-, -f or accurate quantification 1 i- shell form. which is far BETT E R achieved using parameters developed bv Raitg (1561. 19A6). FglmAl .

- r ercductive morphology is similarly conservative, with little

- v..triation in pllial oviduct and bursa copulatrix complex .A.P.-;.rt from size (and absence , of seminal receptacle in F. sancarlosensis) (Figs. 9, .f...i)-=, . Fenial variation is more pronounced, and involves =.,;t'=-.ilv- gitantifigr,ri features that include

17 : number of glandular ridge--; and relative SIZES 1 lobe and filament (Figs. 7, 8, 13, 14, 17, 19, 22, 24. 27, 29, 32, 34,

37).

The relatively high frequency of misclassification cf ungrouped specimens, is partly a result of interpogulation variation within species. Significant variation in shell size and general form is evident in several speciea (Figs. 2d-g, 20g-i, 77,F) and

49 even occurs aMOr. SCMPISE ci F. Monte:UMenSiS from Mcntez.tm Well

proper and ita uPPer outfinw, with classification di the latter only 3. for either =x. cimilar 1 vel cf anatomical variation were not readil parent to us (note only slipht variation in

PENES AMONP SIX POPULATIONS OF F. THOM NI. FIG. 7.7). YET misclassification wA.s =till fr=pu=nt. ThisegThoLJct- reflects occurrence of =untl= variation =tmono ron=P=rifir populations

involvins: comPlex combinations of variables. An adoiticnal

EXPLANATION -- ortr, ufl--.. iE inflated variation within PoPulations due to small samPie sizes

(=-11). Repardlesm of it cits=. Existence of such variation

=;TPCI==T= that di=rriminnt ===:ttirm= ta==d on det from sinPla

PCPULATIONS ( T-POTVP==) for Each sPecies may NOT BE ABLE TO rorrrtly classify additional populations.

50 LITERATURE CITED

BEDUAERT, J.C. and W.E. MILLER. 1973. The molluskS of the arid c-outhwest, with an Arizr, na check list. The University of Arizona

Press, Tucson. AZ, USA. xvi + 271 pp.

, HERSHLER. R. 1385. Ifv mx:tic revision of the Hydrobiidae

(Gaetropoda: Rissoacea) cf. the CuAtrc Cieneqes BSEiri. Coahuila.

7 Mexico. MalA.colociA, 71-1 3,

HERSHLER. R. and F. C-. ThomPson. In Press, North American

Hvdrobiid...f=. (ri.=.strsPrnda: P e area;: Rc...,cc riPticn and systematic relationships of Trvonia StimPscr. 1665 and Fvrpulopsis Call and

Filsbry, 1SSi6. Nautilus: 101.

,. KOHN, A.J. and A.7. R:SES. 1375. Morphcmetry of the ronu..

, shell. Svt-,mAti_ 7r-Cicy,

LANDYE, J.J. 1977. Status of inland aquatic and sami-azuatic mollusks of the American Soutnwest. Submitted to U.S. Department

, , - of Interior, Bureau of 7.-PrIrt Fi=h =ri ===. and Wilf Llife=. Office of

Rare and Endanqsred Spec ice, Vi shingtcri, D.F. 60 pm.

- , - LANDIE. J. J. 1981. Current -.1 t.1.tu--; of P.ndanpered. threatened, and/cr- 1.- re Mollusks of New Mexico and Arizona. Submit tad to

U.S. Dep,=trtment of Interior. Bureau of Sport Fisheries and

Wildlife, Office of Rare •arid End,=tnqered Species, Albuquerque, New

Mexico. Contract N. 14-16-00(D2-80-227. pp.

MAYR. E. 1963. Animal species and evolution. Belknap Press,

Harvard University Press, Cambridge, MA, USA. xiv + 797 pp.

FILSBRY, H.A. 1916. New species of Amnicola from New MEXiC0 and

Utah. Nautilus, 29: 111-112.

FILSERT. H.A. anc J.H. FESRISS. 1909.- Mollu=m= thE

Southwestern States. III: The Huachuca Mcunt=kins. Arizona.

Frohedings of the Academy of Natural Scienoes of FhiledelPhia

(1509. ). pp. 495-516.

TAYLOR, D.W. 1766, A rm=trk=ble snail fauna from Coahuila,

Mexico. V=liger. 5: 152-225.

TAYLCF. D. J. 1'570a. 4. Western freshwater mollusks i-sditor'= summary). P. A.H. Clarke .:ed.). Prmo==inc= hf the

Amrican Malahhlogiczi Union Svmmosi!tm on .=r' and Fnoangered

Malacslogia, 10.

TAYLOR. D.W. 1570b. W==t American Freshwater Molluscs. i:

Bibliography Plet=toc=n= and R=n=nt Diego

Society of Natural Hi=1-,nr,, 4 PP ,

TAY! 7,R. n. W. 1575. and kinlihgraPny- - - - Cenozoic

Freshwater Mollusca of Western North America. MUSEUM C -F

PE1E0MtOlOgy. University of Michigan, Papers on Paleontology Na.

10. 324 Pp.

TAYLOR, D.W. 1583. Status Invest gatihn of Moll;t=!

Mexico Department of Game and Fish. contract Nos. 515-65-01 and

519-65-01-A. Si pp.

United States D=partment of Interior. 1964. Endangered and

threatened wildlife and plants: review of invertebrate wildlife

for listinc as endangered or threatened =',7, ... ,-. 1.=c4. Feder a l

REgi=ter, 45: 21664-21677.

WILET. E.O. 1721. Phvlog ,=netic=. the theory and practice of

phvlogenetic systematics. John Wile. and Sons. New \tor!, N. XV

+ 439 PP. - WILLIAMS, J.E., BOWMAN, J.E. EROT1KS. A.A. ECHELLE, R.J. EDWARDS, D.A. HENDRICKSON, and J.J. LANDYE. Endancoer=d aquatic erosysti=ms in North American deserts with a list of vaniEhins fishes of the region. Journal of the Arizona-Nevada

Academy of Scienrs, Table 1. List of populations receiving morphometric study of shell and anatomy. USNM collection numbers are given for localities. Type localities are indicated with asterisks.

Species and locality Shell Anatomy

1. P. deserta *847202 847206

2. P. bacchus *847203

3. P. conicus *847237 847201

4. P. morrisoni *847231 847220

5. P. montezumensis *847233 847230

6. P. solus *850290

7. P. glandulosus *847205

8. P. simplex —*847236 847222

9. P. confluentis *847234 847229

10. P. sancarlosensis *847226 847217 847224

11. P. thompsoni *847238 847227 847221 847228

12. P. cochisi *847216 Table 2. Generalized cusp formulae for the four tooth types.

Tooth Type

Inner Outer Species Central Lateral marginal marginal

Pyrgulopsis glandulosus 5-1-5 3-1-3(4) 23 31 1-1

P. deserta 4-1-4 (5) 3(4)-1-3 17-19 19-21 1-1

P. bacchus 3(4,5)-1-3(4,5) 2(3)-1-2 18-19 30-31 1-1

P. conicus 4-1-4 3(4)-1-3 19 25-28 1-1

P. morrisoni 6(7)-1-6(5) 3-1-3(4) 19 25-38 1-1

P. montezumensis 8-1-8 3(4,5)-1-5(6) 27-31 28-30 1-1

P. simplex 5-1-5 3-1-5 23-24 28-30 1-1

P. solus 5(6)-1-5(6) 3(4,5)-1-4(5) 21 25-26 1-1

P. confluentis 4-1-5 3-1-4 24-32 23 1-1

P. sancarlosensis 3(5)-1-3(5) 2(3)-1-3 16-22 23-25 1-1

P. cochisi . 4-1-4 3-1-3 22-23 22-23 1-1

P. thompsoni 4(5,6)-1-5(6) 3-1-3(4) 20-24 26-32 1-1

Tryonia gilae 5(6)-1-5(6) 4(5,6)-1-5(6) 24-30 28-31 2-2

T. quitobaquitae 6-1-6 3-1-4 23 26 1-1 Table 3. Results of stepwise discriminant function analysis on thirteen shell

variables.

Standardized weights and correlation of variables

with the first two discriminant functions

Males Females

Variable Function 1 Function 2 Function 1 Function 2

SR 0.56 -0.61 2.44 0.53 -2.21 0.51 2.59 -0.13

SW 2.38 -0.72 1.10 0.27 1.12 0.82 -0.65 0.15

AR 3.91 -0.69 5.54 0.36 -0.91 -3.57 0.10 0.04

AW -0.21 -0.73 0.46 0.37 -0.08 0.68 0.41 0.10

LBW -6.93 -0.69 -5.21 0.48 1.38 0.72 2.57 -0.10

WBW -0.62 -0.79 0.01 0.40 0.89 0.82 0.46 0.03

CD -0.69 -0.30 -0.86 0.38 -1.17 0.31 -0.36 0.01

AB 0.55 -0.55 1.11 0.50 0.64 0.55 -0.71 -0.20

NW -0.12 -0.36 0.27 0.48 0.13 0.33 -0.15 -0.32

SR/SW 1.83 0.06 1.35 0.61 0.56 0.08 -1.44 -0.43

AR/SR -2.52 0.01 -3.28 -0.54 0.58 -0.15 2.98 0.43

LBW/SH 2.61 0.09 1.59 -0.46 -1.08 -0.16 -1.58 0.26

CD/AB 0.64 0.08 1.04 -0.10 0.83 0.01 0.45 0.17 Table 4. Results of stepwise discriminant function analysis on eleven

male anatomical variables.

Standardized weights and correlation of

variables with the first two

discriminant functions

Variable Function 1 Function 2

CTW 0.10 0.50 0.21 0.32

TS/BL -0.04 -0.08 0.78 0.44

L2 0.48 0.44 -0.88 -0.04

BL 0.17 0.03 0.81 0.65

L3/L2 -0.58 -0.06 -0.85 -0.05

GLR 0.87 0.68 0.05 0.03

LFI -0.21 -0.03 -0.49 -0.25 WFI -0.46 0.05 0.20 0.04

CTL -0.05 0.12 -0.31 0.37

LPR/BL 0.56 0.16 -0.06 -0.30

L3 0.28 0.28 1.24 -0.04 Table 5. Results of stepwise discriminant function analysis of thirteen

female anatomical variables.

Standardized weights and correlation

of variables with the first two

discriminant functions

Variable Function 1 Function 2

CTL -0.59 0.49 0.40 0.14

BAG -0.26 0.12 -0.19 -0.40

BUW 0.16 0.23 -0.70 -0.29

COP -0.28 0.15 0.73 0.31

CGW 0.26 0.33 0.14 0.03

Fl 0.25 0.39 0.19 0.17

AGL 0.49 0.40 -0.27 0.09

OV 0.29 0.26 -0.29 -0.09

SRL/BUL -0.07 -0.01 0.70 0.52

OSL 1.01 0.29 -0.19 0.16

BDW 0.62 0.36 0.34 0.03

OCT 0.56 0.38 -0.11 0.01

OSL/CTL -1.29 -0.11 0.70 0.08 Table 6. Classification results of discriminant function analyses. Grouped cases represent topotypes for the 12 species.

Per cent of specimens correctly classified

Data Set Grouped cases

Male shell 89 (n=113)

Female shell 93 (108)

Male anatomy 100 (60)

Female anatomy 100 (60) 1

Appendix 1. Collection localities. Locality data includes appropriate

information from USGS topographic quadrangle sheets (15 minute

series). Numbers in parentheses after locality names refer to

those in Fig. 1.

UNITED STATES

UTAH. WASHINGTON COUNTY: spring on Old Bastion Ranch (1), Hurricane quadrangle, 0.05 km E from NE corner Sec. 13, T. 42S, R. 15W; small spring,

70 in N of interstate 15 (2), St. George, 0.1 km N, 0.1 km E from SW corner

Sec. 15, T. 42S, R. 15W; Green Spring (3), St. George, 0.6 km N, 0.7 km W from

SE corner Sec. 15, T. 42S, R. 15W; spring in Middleton (4), St. George, 0.2 km

W from NE corner Sec. 21, T. 42S, R. 15W; spring on Red Hill (5), St. George,

0.7 km N, 0.6 km E from SW corner Sec. 19, T. 42S, R. 15W; spring on west side of Middleton (6), St. George, 0.1 km S, 0.5 km E from NW corner Sec. 21, T.

42S, R. 15W; Sam Adair Spring (7), St. George, 0.8 km S, 0.6 km E from NW corner Sec. 14, T. 42S, R. 15W. 2

ARIZONA. MOHAVE. Spring just E of Littlefield (8), Littlefield, 0.05 km N,

0.3 km W from SE corner Sec. 25, T. 42N, R. 15W; spring upstream on Virgin

River from Littlefield (9), Littlefield, 1.0 km S, 0.5 km E from NW corner

Sec. 3, T. 40 N, R. 15W; spring NW of Littlefield (10), Littlefield, 1.0 km S,

0.5 km W from NW corner Sec. 3, T. 40N, R. 15W; spring NW of Littlefield (10), km S, 0.5 km W from NE corner Sec. 4, T. 40N, R. 15E; Grapevine Spring (11),

Gypsum Hills, 0.4 km W from NE corner Sec. 26, T. 34N, R. 16W; Burns Spring

(12), Burns Spring, 0.1 km S, 0.2 km W from NE corner Sec. 9, T. 22N, R. 20W;

Dripping Springs (13), Mount Nutt, 0.6 km S, 0.7 km W from NE corner Sec. 4,

T. 19N, R. 19W; Cool Spring (14), Mount Nutt, 0.5 km N, 0.5 km W from SE

corner Sec. 20, T. 19N, R. 19W.

YAVAPAI. Tavasci Springs (15), Clarkdale, 0.4 km N, 0.4 km E from SW corner

Sec. 15, T. 16N, R. 3E; Page Springs (16), Page Springs, 0.1 km N, 0.3 km W

from SE corner Sec. 23, T. 16N, R. 4E; Montezuma Well (17), Montezuma Well,

0.9 km S, 1.1 km E from NW corner Sec. 31, T. 15N, R. 6E; Brown Spring (18),

Horner Mountain, 0.3 km N, 0.4 km W from SE corner of Sec. 23, T. 12N, R. 9E;

Fossil Springs (19), Strawberry, 0.4 km N, 0.2 km W from SE corner Sec. 14,

T. 12N, R. 7E; Nelson Place Spring (21), Tule Mesa, 6.0 km S, 4.3 km E from NW

corner of quadrangle; spring 150 m E of Nelson Place Spring (22), Tule Mesa,

6.1 km S, 4.8 km E from NW corner of Sec. 15, T. 11N, R. 5E.

GILA. Spring near Strawberry (20), Strawberry, 0.3 km N, 0.3 km E from SW

corner Sec. 22, T. 12N, R. 7E.

APACHE. Spring on N side of Blanket Creek at Three Forks (23), Big Lake, 0.5

km N, 0.2 km W from SE corner Sec. 6, T. 5N, R. 29E; spring at Three Forks

(24), Big Lake, 0.2 km N from SE corner Sec. 6, T. 5N, R. 29E. 3

GRAHAM. Small spring near Bylas (25), Bylas, 0.2 km N, 0.3 km E from SW

corner Sec. 17, T. 3S, R. 22E; spring near Gila River bridge NW of Bylas (26),

Bylas, 0.1 km N, 0.3 km E from SW corner Sec. 17, T. 3S, R. 22E; large spring

N of Bylas (27), Bylas, 0.1 km N, 0.3 km E from SW corner Sec. 28, T. 3S, R.

22E; springs W of Tom Niece Springs (28), Fort Thomas, 0.8 km S from NE corner

Sec. 21, T. 4S, R. 23E; seeps W of Tom Niece Springs (29), Fort Thomas, 0.6 km

S, 0.4 km W from SE corner Sec. 21, T. 4S, R. 23E; Cold Springs (30), Fort

Thomas, 0.1 km N, 0.5 km W from SE corner Sec. 8, T. 5S, R. 24E.

PIMA. Quitobaquito Springs (31), Quitobaquito, 6.8 km S. 1.8 km W from NE

corner of quadrangle, T. 17S, R. 7W.

SANTA CRUZ. Cottonwood Springs (32), Sonoita, 0.6 km S. 0.1 km W from NE

corner Sec. 33, T. 20S, R. 16E; Monkey Spring (33), Sonoita, 0.5 km N, 0.4 km

E from SW corner Sec. 3, T. 21S, R. 16E; Canelo Hills Cienega (34), O'Donnell

Canyon, 0.6 km N, 0.1 km W from SE corner Sec. 32, T. 21S, R. 18E; Sheehy

Spring (35), Lochiel, NW corner Sec. 12, T. 24S, R. 17E; Peterson Ranch

Springs (36), Huachuca Peak, 0.5 km N, 0.6 km W from SE corner Sec. 3, T. 23S,

R. 19E.

COCHISE. Springs at San Bernardino Ranch (38), College Peaks, 0.3 km N, 0.8

km E from SE corner Sec. 15, T. 34S, R. 30E.

MEXICO

SONORA. Ojo Caliente, 44 km S of Nogales (37). Appendix 2. Shell data. (IS M catalog numbers are given for localities. Sample sizes are given in parentheses (fmales, /females).

Asterisked localities have significant (two tailed t-test, p<.05) sexual dimorphism in shell height (SH). Measurements

are in mm.

Parameters

SH SW AM AW LBW WHW CD AS NW

Species and locality

P. glandulosus *847205 x 2.24 2.56 1.67 1.93 1.13 1.28 1.03 1.12 1.84 2.13 1.42 1.60 0.18 0.19 0.68 0.78 3.78 3.89 (a = 8,7) • 0.18 0.15 0.13 0.08 0.09 0.10 0.10 0.06 0.14 0.18 0.13 0.10 0.05 0.01 0.07 0.05 0.16 0.13

P. deserts *847202 ; 1.51 1.41 1.28 1.20 0.85 0.80 0.75 0.74 1.30 1.24 1.05 0.95 0.09 0.12 0.41 0.40 3.44 3.28 (n = 8,9) s 0.17 0.13 0.12 0.10 0.06 0.06 0.05 0.06 0.11 0.12 0.10 0.07 0.01 0.03 0.04 0.05 0.18 0.20

847206 x 1.47 1.76 1.15 1.36 0.74 0.87 0.71 0.82 1.22 1.41 0.99 1.16 0.13 0.12 0.42 0.50 3.47 3.69 (n = 8) • 0.13 0.15 0.06 0.04 0.04 0.03 0.04 0.05 0.08 0.09 0.04 0.06 0.10 0.02 0.04 0.06 0.16 0.29

P. bacchus 84203 x 2.57 2.54 1.79 1.84 1.23 1.23 1.13 1.14 2.02 2.02 1.58 1.58 0.20 0.18 0.78 0.74 4.18 4.15 (n 10) s 0.23 0.16 0.08 0.07 0.05 0.05 0.05 0.07 0.12 0.11 0.08 0.06 0.03 0.03 0.11 0.06 0.12 0.18

F. conicus - 847237 x 2.36 2.31 1.54 1.57 1.11 1.11 0.96 0.96 1.83 1.81 1.29 1.33 0.13 0.15 0.68 0.66 4.11 4.09 (n = 9,8) • 0.17 0.14 0.11 0.09 0.07 0.08 0.07 0.06 0.12 0.10 0.08 0.09 0.02 0.02 0.07 0.04 0.13 0.13

*847201 it 2.17 2.47 1.43 1.65 1.01 1.11 0.84 0.98 1.66 1.88 1.23 1.37 0.13 0.15 0.61 0.69 4.07 4.25 (n + 7,9) • 0.21 0.13 0.14 0.10 0.09 0.07 0.08 0.05 0.12 0.06 0.10 0.05 0.01 0.02 0.04 0.04 0.31 0.13 P. morrisoni *847231 ; 1.89 2.08 1.43 1.54 0.92 0.98 0.83 0.91 1.49 1.63 1.14 1.26 0.11 0.12 0.56 0.59 3.78 4.00 (n = 9) s 0.10 0.14 0.07 0.11 0.07 0.08 0.05 0.07 0.10 0.10 0.05 0.08 0.02 0.02 0.03 0.05 0.15 0.13 847220 x 2.29 2.37 1.64 1.68 1.11 1.12 0.97 0.99 1.80 1.82 1.39 1.43 0.15 0.14 0.67 0.66 4.13 4.03 (n + 8,10) • 0.19 0.18 0.10 0.11 0.10 0.07 0.08 0.06 0.14 0.12 0.08 0.08 0.02 0.01 0.07 0.06 0.19 0.22 Appendix 2 Cont'd.

P. montezumensis 847233 i 2.12 2.18 1.41 1.51 0.95 0.99 0.87 0.93 1.55 1.65 1.25 1.32 0.15 0.17 0.59 0.62 4.00 4.06 (n * 10,8) 0.28 0.15 0.20 0.09 0.14 0.05 0.10 0.08 0.19 0.09 0.14 0.08 0.03 0.02 0.05 0.05 0.20 0.26

*847230 i 2.06 2.41 1.42 1.66 0.93 1.07 0.81 0.95 1.54 1.74 1.17 1.40 0.11 0.15 0.56 0.65 4.05 4.14 (n •. 10,9) s 0.24 0.14 0.12 0.06 0.07 0.06 0.07 0.04 0.19 0.08 0.12 0.09 0.02 0.02 0.06 0.65 0.37 0.13 P. simplex *847236 i 2.08 2.31 1.43 1.55 1.01 1.08 0.87 0.94 1.67 1.80 1.18 1.27 0.13 0.14 0.59 0.63 3.81 4.00 (n 9,10) 0.12 0.09 0.11 0.07 0.07 0.07 0.06 0.06 0.09 0.10 0.08 0.08 0.01 0.02 0.04 0.05 0.21 0.20

847222 i 2.18 2.21 1.42 1.56 0.97 1.05 0.87 0.90 1.65 1.68 1.25 1.26 0.13 0.13 0.63 0.67 4.03 4.00 (n * 10,9) 8 0.19 0.16 0.11 0.11 0.09 0.11 0.007 0.06 0.15 0.08 0.08 0.06 0.01 0.02 0.07 0.11 0.18 0.18 P. solus - *850290 i 1.57 1.72 1.17 1.29 0.78 0.87 0.73 0.82 1.29 1.45 1.00 1.08 0.11 0.11 0.46 0.50 3.50 3.73 (n * 8,10) 8 0.08 0.16 0.04 0.12 0.03 0.08 0.03 0.07 0.05 0.14 0.03 0.08 0.02 0.02 0.04 0.05 0.13 0.18 P. confluentis 847234 i 2.81 3.01 1.80 1.92 1.18 1.24 1.10 1.16 2.09 2.18 1.65 1.76 0.19 0.24 0.82 0.85 4.30 4.35 (n = 10) 8 0.29 0.68 0.16 0.15 0.10 0.10 0.11 0.09 0.18 0.19 0.14 0.18 0.03 0.07 0.09 0.14 0.20 0.32

*847229 i 1.62 1.83 1.30 1.42 0.83 0.93 0.77 0.84 1.36 1.50 1.08 1.21 0.14 0.15 0.50 0.53 3.73 3.68 (n = 10) 8 0.12 0.80 0.09 0.06 0.06 0.04 0.04 0.03 0.09 0.10 0.08 0.06 0.02 0.03 0.05 0.04 0.14 0.21 P. sancarlosensis 847226 i 1.90 2.01 1.28 1.33 0.92 0.93 0.82 0.85 1.56 1.66 1.09 1.15 0.15 0.15 0.56 0.62 3.90 4.06 (n ■ 13) • 0.19 0.20 0.08 0.11 0.07 0.06 0.05 0.04 0.14 0.90 0.08 0.07 0.02 0.03 0.07 0.06 0.26 0.17 847217 i 1.34 1.54 1.06 1.20 0.74 0.83 0.67 0.75 1.17 1.34 0.83 0.67 0.08 0.11 0.40 0.46 3.40 3.65 10) • 0.17 0.34 0.10 0.16 0.07 0.11 0.06 0.11 0.12 0.21 0.10 0.15 0.01 0.02 0.05 0.11 0.21 0.27 847224 1 1.50 1.62 1.11 1.20 0.75 0.82 0.73 0.77 1.28 1.40 0.95 1.02 0.11 0.13 0.45 0.48 3.33 3.53 (n * 9) • 0.06 0.82 0.04 0.09 0.03 0.05 0.03 0.05 0.05 0.11 0.03 0.08 0.02 0.03 0.03 0.07 0.13 0.36 Appendix 2 Cont'd.

P. cochisi *847218 i 1.41 1.54 0.89 1.10 0.63 0.69 0.57 0.63 1.11 1.21 0.81 0.88 0.11 1.12 0.43 0.48 3.56 3.69 9) s 0.09 0.12 0.05 0.09 0.03 0.05 0.03 0.05 0.05 0.08 0.04 0.05 0.01 0.02 0.03 0.04 0.21 0.17 P. thoupsoni -*847238 i 2.67 2.41 1.68 1.59 1.20 1.12 1.05 1.00 2.03 1.90 1.43 1.42 0.18 0.14 0.77 0.70 4.53 4.14 (n 10,9) s 0.24 0.17 0.11 0.09 0.08 0.08 0.07 0.07 0.13 0.12 0.08 0.11 0.02 0.02 0.06 0.05 0.18 0.13 847221 i 1.97 2.08 1.43 1.43 0.97 0.97 0.87 0.87 1.60 1.59 1.18 1.18 0.12 0.13 0.58 0.59 4.03 4.03 (n.8,9) s 0.08 0.18 0.06 0.12 0.04 0.06 0.04 0.05 0.08 0.10 0.05 0.08 0.01 0.01 0.02 0.04 0.21 0.20 *847228 i 2.15 2.51 1.40 1.62 0.96 1.12 0.90 1.01 1.64 1.89 1.29 1.47 0.14 0.15 0.62 0.73 4.05 4.35 (n . 10) s 0.25 0.31 0.10 0.11 0.09 0.08 0.07 0.08 0.15 0.17 0.10 0.10 0.02 0.01 0.07 0.07 0.23 0.29 847227 i 1.91 1.85 1.19 1.23 0.83 0.87 0.75 0.76 1.43 1.47 1.09 1.13 0.14 0.14 0.54 0.57 3.91 3.89 (n 8,9) s 0.19 0.19 0.08 0.05 0.06 0.04 0.07 0.04 0.12 0.07 0.07 0.06 0.04 0.02 0.05 0.05 0.38 0.18 Appendix 3. Female anatomical data. USNM catalog numbers are given for the localities. For all samples, n •. 5. Measurements are in mm.

Parameters

Species and locality BL OV OSL OSW OCT CTL CTW F1 CGL CGW AGL AGW COL COP COD BUL BOW BOW BAG SRI. SRW P. glandulosus 847205 x 4.22 0.72 0.24 0.29 0.22 1.00 0.20 18.80 0.59 0.29 0.62 0.38 0.27 0.20 0.09 0.27 0.18 0.05 0.18 0.26 0.08 s 0.36 0.09 0.04 0.44 0.03 0.07 0.19 1.30 0.06 0.02 0.07 0.02 0.08 0.02 0.02 0.02 0.05 0.01 0.04 0.07 0.02 P. deserta 847202 2.70 0.58 0.16 0.06 0.18 0.70 0.17 12.40 0.39 0.27 0.50 0.33 0.18 0.13 0.06 0.42 0.22 0.05 0.30 0.16 0.07 m 0.26 0.21 0.04 0.01 0.04 0.05 0.02 0.90 0.02 0.05 0.09 0.04 0.02 0.04 0.01 0.07 0.03 0.01 0.05 0.04 0.01 P. bacchus 847203 x 6.07 0.97 0.28 0.08 0.31 1.15 0.25 17.40 0.62 0.37 0.82 0.44 0.35 0.12 0.11 0.39 0.29 0.12 0.25 0.16 0.07 a 0.55 0.08 0.02 0.01 0.02 0.05 0.05 1.67 0.05 0.02 0.06 0.04 0.06 0.04 0.02 0.06 0.04 0.01 0.06 0.05 0.02 P. conicus 847237 x 5.07 0.89 0.29 0.06 0.30 1.10 0.23 20.00 0.57 0.30 0.70 0.38 0.29 0.16 0.09 0.29 0.18 0.08 0.12 0.17 0.09 a 0.52 0.13 0.02 0.00 0.02 0.07 0.03 1.00 0.05 0.03 0.10 0.04 0.03 0.03 0.01 0.04 0.20 0.01 0.04 0.03 0.02 P. morrisoni 847231 3.78 0.74 0.25 0.09 0.20 0.82 0.18 16.40 0.49 0.27 0.47 0.32 0.24 0.07 0.09 0.24 0.16 0.08 0.17 0.20 0.06 a 0.14 0.20 0.02 0.01 0.04 0.14 0.02 1.14 0.06 0.02 0.07 0.05 0.01 0.06 0.01 0.04 0.03 0.01 0.04 0.05 0.00 847220 5.36 0.89 0.36 0.10 0.25 1.21 0.25 18.20 0.67 0.41 0.86 0.52 0.24 0.25 0.09 0.24 0.18 0.08 0.12 0.26 0.10 s 0.41 0.11 0.06 0.01 0.05 0.17 0.03 0.84 0.08 0.01 0.12 0.09 0.05 0.05 0.01 0.03 0.03 0.02 0.06 0.04 0.02 P. montezusensis 847233 x 4.63 0.60 0.26 0.10 0.26 1.06 0.20 17.40 0.60 0.34 0.68 0.41 0.27 0.20 0.10 0.33 0.25 0.07 0.22 0.21 0.07 • 0.16 0.08 0.04 0.02 0.03 0.02 0.02 1.52 0.03 0.04 0.04 0.04 0.04 0.03 0.01 0.05 0.04 0.01 0.04 0.03 0.02 P. simplex -. 847236 i 3.87 0.45 0.21 0.09 0.24 1.03 0.16 17.60 0.49 0.23 0.54 0.30 0.22 0.10 0.08 0.29 0.14 0.07 0.15 0.17 0.05 • 0.32 0.22 0.02 0.03 0.06 0.09 0.01 0.14 0.08 0.02 0.10 0.03 0.02 0.02 0.00 0.12 0.01 0.01 0.05 0.04 0.01 Appendix 3 Contd.

-P. solus 850290 ; 3.44 0.51 0.19 0.07 0.18 0.78 0.17 12.60 0.46 0.27 0.50 0.32 0.22 0.12 0.07 0.18 0.17 0.05 0.10 0.15 0.06 s 0.23 0.08 0.02 0.01 0.03 0.07 0.01 0.89 0.04 0.04 0.06 0.03 0.03 0.01 0.01 0.01 0.04 0.01 0.00 0.02 0.00 P. confluentis 847234 x 6.15 0.91 0.33 0.06 0.30 1.43 0.20 20.60 0.82 0.40 0.87 0.47 0.34 0.19 0.08 0.39 0.27 0.08 0.25 0.37 0.11 s 0.56 0.13 0.02 0.02 0.03 0.20 0.04 1.52 0.03 0.03 0.06 0.05 0.07 0.02 0.01 0.04 0.04 0.01 0.04 0.07 0.02 P. sarcarlosensis 847226 x 3.96 0.85 0.23 0.06 0.22 0.92 0.19 17.80 0.54 0.26 0.49 0.37 0.40 0.01 0.09 0.49 0.26 0.06 0.29 0.00 0.00 s 0.18 0.14 0.05 0.02 0.01 0.01 0.01 1.30 0.06 0.02 0.01 0.04 0.04 0.01 0.01 0.05 0.03 0.02 0.02 0.00 0.00 847217 x 2.66 0.41 0.22 0.07 0.19 0.73 0.17 17.20 0.37 0.23 0.37 0.28 0.30 0.00 0.10 0.22 0.18 0.07 0.17 0.00 0.00 0.26 0.03 0.05 0.02 0.03 0.09 0.02 1.79 0.05 0.02 0.03 0.02 0.02 0.00 0.02 0.03 0.02 0.01 0.02 0.00 0.00 P. cochisi 847218 x 2.69 0.39 0.19 0.55 0.17 0.62 0.13 14.20 0.44 0.20 0.41 0.26 0.14 0.05 0.05 0.17 0.15 0.04 0.11 0.11 0.02 s 0.27 0.04 0.01 0.02 0.03 0.05 0.03 0.84 0.02 0.02 0.06 0.02 0.02 0.02 0.01 0.03 0.02 0.01 0.03 0.01 0.01

-P. thompsoni 847238 i 5.19 0.72 0.19 0.07 0.32 1.07 0.22 18.20 0.68 0.33 0.73 0.40 0.31 0.13 0.07 0.32 0.20 0.07 0.26 0.23 0.06 s 0.94 0.14 0.03 0.03 0.02 0.13 0.02 1.30 0.16 0.06 0.15 0.05 0.06 0.07 0.01 0.06 0.03 0.02 0.06 0.06 0.01 847227 i 5.67 0.68 0.33 0.10 0.38 0.31 0.26 19.80 0.63 0.34 0.69 0.44 0.22 0.17 0.07 0.31 0.15 0.05 0.27 0.16 0.06 s 0.36 0.11 0.03 0.02 0.04 0.11 0.003 1.30 0.06 0.04 0.10 0.05 0.04 0.04 0.01 0.05 0.04 0.02 0.06 0.02 0.01 Appendix 4. Hale anatomical data. OSNM catalog numbers are given for the localities. For all samples, n ■ 5.

Measurements are in mm.

Parameters

Species and Locality BL TS OSL OSW OCT CTL CTW FI LPR WPR L2 L3 PW LFI WFI. GLR

P. montezumensis 847233 3.92 1.82 0.28 0.09 0.21 0.99 0.23 15.80 0.59 0.29 1.09 0.92 0.49 0.32 0.13 3.60 0.26 0.40 0.04 0.02 0.02 0.18 0.02 0.84 0.04 0.03 0.12 0.13 0.02 0.07 0.01 0.55

P. simplex 847236 3.14 0.89 0.22 0.09 0.21 0.93 0.16 17.40 0.37 0.22 0.67 0.52 0.25 0.38 0.10 1.00 8 0.80 0.32 0.05 0.02 0.03 0.04 0.03 1.14 0.04 0.04 0.08 0.13 0.04 0.12 0.01 0.00

P. Bolus 850290 i 3.14 1.38 0.20 0.07 0.17 0.72 0.14 12.60 0.34 0.20 0.40 0.27 0.23 0.18 0.07 2.00 s 0.23 0.23 0.03 0.01 0.03 0.09 0.02 1.14 0.03 0.01 0.05 0.09 0.03 0.03 0.01 0.00

P. confluentis 847234 i 5.52 2.53 0.35 0.08 0.26 1.24 0.21 19.40 0.55 0.31 1.24 0.83 0.49 0.33 0.15 5.40 s 0.38 0.28 0.02 0.03 0.03 0.08 0.06 1.82 0.06 0.02 0.14 0.12 0.09 0.04 0.07 0.55 P. sarcarloensis 847226 i 3.34 0.89 0.23 0.07 0.20 0.96 0.18 18.00 0.56 0.26 1.09 0.78 0.44 0.72 0.19 1.80 s 0.32 0.22 0.04 0.01 0.03 0.06 0.02 2.00 0.10 0.05 0.12 0.11 0.03 0.08 0.03 0.84 847217 i 2.64 0.71 0.24 0.06 0.14 0.74 0.15 14.00 0.46 0.20 0.67 0.56 0.34 0.56 0.16 2.00 a 0.41 0.07 0.03 0.02 0.03 0.13 0.02 1.58 0.04 0.03 0.09 0.12 0.07 0.06 0.02 0.00 P. glanuloslus 847205 i 3.44 0.88 0.31 0.11 0.18 0.96 0.20 17.40 0.56 0.26 1.42 1.10 0.43 0.36 0.11 4.80 s 0.32 0.04 0.04 0.02 0.02 0.04 0.03 0.55 0.04 0.04 0.21 0.20 0.07 0.08 0.02 0.45 P. deserta 847202 i 2.44 0.75 0.18 0.07 0.15 0.55 0.18 11.20 0.48 0.21 0.53 0.45 0.34 0.39 0.16 3.00 s 0.14 0.13 0.03 0.001 0.03 0.09 0.02 0.84 0.04 0.05 0.04 0.04 0.03 0.03 0.02 0.00 Appendix 4 Cont'd.

P. bacchus 847203 6.06 3.29 0.31 0.09 0.29 1.17 0.30 17.80 0.81 0.39 0.77 0.64 0.35 0.32 0.17 2.00 0.25 0.38 0.05 0.02 0.03 0.10 0.03 1.92 0.15 0.06 0.15 0.17 0.02 0.07 0.03 0.71 P. conicus 847237 i 4.68 1.92 0.26 0.08 0.24 1.04 0.25 18.60 0.47 0.28 0.69 0.49 0.36 0.34 0.12 1.00 0.53 0.36 0.06 0.02 0.01 0.12 0.04 1.14 0.07 0.04 0.20 0.21 0.03 0.05 0.01 0.00 P. morrisoni 847231 i 3.45 1.60 0.24 0.09 0.16 0.69 0.17 16.20 0.38 0.22 1.02 0.72 0.37 0.30 0.09 1.00 0.17 0.24 0.04 0.02 0.02 0.07 0.01 0.84 0.07 0.03 0.12 0.18 0.06 0.04 0.01 0.00 847220 4.69 2.54 0.31 0.10 0.26 0.97 0.23 17.20 0.56 0.31 1.11 0.85 0.44 0.53 0.14 1.00 0.57 0.26 0.03 0.02 0.02 0.03 0.03 2.39 0.08 0.03 0.13 0.17 0.05 0.07 0.01 0.00 P. cochisi 847218 2.50 0.94 0.17 0.06 0.15 0.58 0.12 13.40 0.28 0.17 0.45 0.45 0.23 0.27 0.09 2.00 s 0.24 0.05 0.01 0.01 0.02 0.04 0.02 1.34 0.01 0.03 0.03 0.03 0.02 0.03 0.01 0.00 P. thompsoni 847238 i 5.47 2.96 0.20 0.08 0.29 1.12 0.23 19.00 0.37 0.26 0.47 0.41 0.42 0.46 0.15 1.00 s 0.89 0.39 0.04 0.01 0.05 0.15 0.04 0.71 0.10 0.04 0.11 0.11 0.08 0.06 0.02 0.00 847221 x 3.80 2.01 0.26 0.07 0.20 0.93 0.19 16.40 0.3 0.22 0.69 0.56 0.32 0.27 0.08 1.00 0.27 0.31 0.02 0.02 0.04 0.09 0.02 0.90 0.03 0.04 0.08 0.10 0.06 0.05 0.02 0.00 847228 x 5.12 2.14 0.30 0.09 0.33 1.14 0.24 17.60 0.34 0.25 0.50 0.46 0.28 0.39 0.10 1.00 s 0.58 0.19 0.07 0.01 0.04 0.20 0.06 1.52 0.03 0.03 0.06 0.05 0.09 0.05 0.02 0.00 847227 i 3.64 1.35 0.24 0.08 0.26 0.87 0.19 15.60 0.25 0.17 0.82 0.66 0.28 0.29 0.13 1.00 • 0.35 0.18 0.05 0.01 0.04 0.12 0.01 1.34 0.02 0.03 0.07 0.04 0.04 0.04 0.01 0.00 FIGURE CAPTIONS

FIGURE 1.-Map of Arizona drainage, showing the 38 collecting sites numbered as in Appendix 1.

FIGURE 2.-Pyrgulopsis glandulosus new species and Pyrgulopsis confluentis new species: a-c, P. glandulosus (a. holotype. USNM

859037, Nelson Place Spring, Yavapai County. Arizona; b, c. same l ocality, USNM 847205); d-g, P. confluentis (d, holotype. USNM

859053, spring on N side of Blanket Creek, Apache County,

Arizona; e. same locality. USNM 847234; f. g. USNM 847229, spring at Three Forks, Apache County, Arizona) (bar = 0.75 mm).

FIGURE 3.-Right l ateral aspect of relaxed F. glandulosus new species (without shell), USNM 847205, Nelson Place Spring,

Yavapai County. Arizona: darkly pigmented epithelia appear black. glandular ridges in penis screened, and subepithelial

- pigment Cr.ndicated by groups of stipples (Abbreviations: Ft = foot, He = head, Opl = operculigerous lobe, Pn = penis).

FIGURE 4.-SEM micrographs of penis and cephalic tentacles of P. glandulosus new species, USNM 847205, Nelson Place Spring,

Yavapai County, Arizona: a, close-up of glandular ridge (bar =

120 um); b, dorsal aspect of penis (bar = 0.33 mm); c, right tentacle, showing sparse ciliation (bar = 176 um); d, l eft tentacle (bar = 136 um).

FIGURE 5.-Anatomy of P. glandulosus new species. USNM 847205,

Nelson Place Spring, Yavapai County, Arizona: a, head and e044, contents of pallial cavity, which Nwee--treert exposed by slitting D pallial roof mid-dorsally: b. ctenidium and adjacent structures; right lateral aspect of 4 stomach; d. coiled

1

seminal vesicle, with et-position of the testis indicated by L 1 dashed lines; e, right lateral aspect of elprostate gland, with thickened curving line indicating CL;osterior end of

pallial cavity (Abbreviations: Ast = anterior stomach chamber; Cc

= caecal chamber: Cm = columellar muscle; Ct = ctenidium; Dg0 =

opening of the digestive gland into the stomach; I n = intestine;

Ki = kidney; Mc = mantle collar: Oes = oesophagus; Os

osphradium: Fn = penis: Fr = prostate; Pst = posterior stomach

chamber: Sn = snout; Sts = style sac; Tn = cephalic tentacle: Vd

= vas deferens).

FIGURE 6.-Radula of F. glandulosus new species. USNM 847205,

Nelson Place Spring. Yavapai County, Arizona: a, centrals (bar =

13.6 um); b. l aterals (bar = 15.0 um); c. l aterals and inner

marginals ( bar = 15.0 um): d, outer marginals ( bar = 5.0 um).

FIGURE 7.-Penes of P. glandulosus new species. USNM 847205.

Nelson Place Spring, Yavapai County, Arizona: Dorsal aspects

_

TO l eft, ventral aspects to aitl ° right; screened areas

indicate glandular ridges (Abbreviations: Acc = accessory crest;

Pt = penial filament; Plo = penial l obe).

FIGURE 8.-Penes of P. glandulosus new species: a, b, USNM

847205, Nelson Place Spring, Yavapai County. Arizona; c, USNM

847232, spring 150 in E of Nelson Place Spring, Yavapai County,

Arizona.

FIGURE 9.-Pallial oviducts of Pyrgulopsis spp., viewed from the

left side (two tissue sections indicated by screens): a, P.

glandulosus new species, USNM 847205, Nelson Place Spring,

Yavapai County, Arizona (shows position of other structures at

- anterior end of 0pa1lial cavity, with 4 mantle collar

2 -- indicated by edashed line); b, bursa copulatrix from above; c. P. deserta (Pilsbry), USNM 847202, spring upstream on Virgin

River from Littlefield, Mohave County, Arizona; d. P. bacchus new

species, USNM 847203, Grapevine Spring, Mohave County, Arizona;

F. conicus new species. USNM 847237, Dripping Springs. Mohave

County, Arizona; f. P. morrisoni new species. USNM 847231, Page

Springs. Yavapai County, Arizona; g. P. montezumensus new species, USNM 847233, Montezuma Well, Yavapai County. Arizona

(Abbreviations: Ag = albumen gland; Eu = bursa copulatrix; Cg = capsule gland; Cga = capsule gland opening; Cm = columellar

muscle; Dbu = duct of bursa; In = intestine).

FIGURE 10.-P. deserta (Pilsbryi and P. bacchus new species: a-

d. F. deserta (a, holotype, ANSP 12112, Washington County. Utah;

b, USNM 847210, spring in Middleton, Washington County, Utah; c.

USNM 847239. spring on Red Hill, Washington County. Utah; d. USNM

847202, spring upstream on Virgin River from Littlefield, Mohave

County, Arizona); e-g, P. bacchus (e. USNM 859037, holotype,

Grapevine Spring, Mohave County, Arizona; f, g, USNM 847203, same

locality) (bar = 0.75 mm).

FIGURE 11.-Radula of P. deserta (Filsbry) and P. bacchus new species: a-c, P. deserta, USNM 847202, spring upstream on Virgin

River from Littlefield, Mohave County, Arizona (a, centrals, bar

= 12.0 um; b, l aterals and inner marginals, bar = 15.0 um; c, outer marginals, bar = 17.6 um); d-f, P. bacchus. USNM 847203,

Grapevine Spring, Mohave County, Arizona (d, centrals, bar = 20.0 um; e, centrals, l aterals, inner marginals, bar = 38 um; f, laterals, inner and outer marginals, bar = 38 um).

3 FIGURE 12.-Protoconchs and opercula of Arizona hydrobiids: a, protoconch, P. montezumensus new species, USNM 847233. Montezuma

Well, Yavapai County, Arizona (bar = 136 um); b, protoconch, P. bacchus new species, USNM 847203, Grapevine Spring, Mohave

County, Arizona (bar = 136 um); c, protoconch, Tryonia gilae new species. USNM 847257, small spring near Bylas, Graham County.

Arizona (bar = 75 um); d, operculum, F. montezumensus. same lot as above (bar = 0.33 mm); e. opPrculum. P. sancarlosensis new species, USNM 847226, springs W of Tom Niece Springs, Graham

County, Arizona (bar = 0.27 ram); f, operculum, T. gilae. same lot as above (bar = 176 um).

FIGURE 13.-Penes of P. deserta (Pilsbryi: a.c, USNM 847239, spring on Red Hill, Washington County, Utah; b, USNM 847244. spring on west side of Middleton. Washington County. Utah; d, e.

USNM 847202, spring upstream on Virgin River from Littlefield,

Mohave County, Utah.

FIGURE 14.-Penes of P. bacchus new species. USNM 847203,

Grapevine Spring, Mohave County, Arizona.

FIGURE 15.-P. conicus new species and P. morrisoni new species: a-d, P. conicus (a, holotype, USNM 859039, Dripping Springs,

Mohave County, Arizona; b, USNM 847203, same locality; c, d. USNM

847204, Burns Spring, Mohave County, Arizona); e-h, P. morrisoni

(e, holotype, USNM 859041, Page Springs, Yavapai County, Arizona; f, USNM 847231, same locality; g, h, USNM 847220, Tavasci

Springs, Yavapai County, Arizona) (bar = 0.75 mm).

FIGURE 16.-Radula of P. conicus new species: a, centrals (bar =

12.0 um); b, l aterals (bar = 8.6 um); c, l aterals and inner marginals (bar = 16.2 um); d, outer marginals (bar = 6.0 um).

4 FIGURE 17.-Penes of P. conicus new species: a, USNM 847204,

Burns Spring, Mohave County. Arizona; b-d, USNM 847237. Dripping

Springs, Mohave County, Arizona; e, USNM 847201, Cool Spring,

Mohave County, Arizona.

FIGURE 18.-Radula of P. morrisoni new species: a. centrals,

l aterals, inner marginals (bar = 13.6 um); b, centrals (bar =

12.0 um); c, i nner marginals (bar = 13.6 um); d, inner and outer marginals (bar = 23.1 um).

FIGURE 19.-Penes of P. morrisoni new species: a.b. USNM 847231,

Page Springs, Yavapai County. Arizona; c, d. USNM 847220, Tavasci

Springs, Yavapai County, Arizona.

FIGURE 20.-P. montezumensus new species, P. solus new species, and P. simplex new species: a-c, P. montezumensus (a. USNM

859043, holotype, Montezuma Well, Yavapai County, Arizona; b. c.

USNM 847233, same locality); d-f, P. solus (d. USNM 869045. holotype, Brown Spring, Yavapai County, Arizona; e, f, USNM

850290, same locality); g-j, P. simplex (g, USNM 859049, holotype, spring near Strawberry, Gila County, Arizona; h, USNM

847236, l ocality as above; i, j, USNM 847222, Fossil Springs,

Yavapai County, Arizona) (bar = 0.75 mm).

FIGURE 21.-Radula of P. montezumensus new species, USNM 847233,

Montezuma Well, Yavapai County, Arizona: a, centrals (bar = 12.0 um); b, l aterals (bar = 8.6 um); c, l aterals and inner marginals

(bar = 16.2 um); d, outer marginals (bar = 4.3 um).

FIGURE 22.-Penes of P. montezumensus new species: a. b. USNM

847233, Montezuma Well, Yavapai County, Arizona; c, USNM 847230, locality as above (but from spring outflow).

5 FIGURE 23.-Radula of P. solus new species. USNM 850290. Brown

Spring, Yavapai County, Arizona: a, centrals (bar = 8.6 um); b,

l aterals and inner marginals (bar = 7.5 um); c, inner marginals

(bar = 6.0 um); d, outer marginals (bar = 6.0 um).

FIGURE 24.-Penes of P. solus new species, USNM 850290. Brown

Spring, Yavapai County, Arizona.

FIGURE 25.-Palli3l oviducts of Fyrgulopsis spp.: a. P. solus new species, USNM 850290. Brown Spring, Yavapai County, Arizona; b. P. simplex new species, USNM 847236. spring near Strawberry,

Gila County, Arizona; c. P. confluentis new species, USNM 847234, spring on N side etBlanket Creek at Three Forks, Apache County,

Arizona; d, P. cochisi new species. USNM 847218, Springs at San

Bernardino Ranch, Cochise County. Arizona; e. P. sancarlosensis new species, USNM 847226, springs W of Tom Niece Springs, Graham

County. Arizona; f, P. thompsoni new species, USNM 847238.

Peterson Ranch Springs, Santa Cruz County, Arizona.

FIGURE 26.-Radula of P. simplex new species, USNM 847236, spring near Strawberry, Gila County, Arizona: a, centrals (bar =

12.0 um); b, laterals (bar = 13.6 um); c, inner marginals ( bar =

10.0 um); d, outer marginals ( bar = 7.5 um).

FIGURE 27.-Penes of P. simplex new species: a, b, USNM 847236. spring near Strawberry, Gila County, Arizona; c, USNM 847222,

Fossil Springs, Yavapai County, Arizona.

FIGURE 28.-Radula of P. confluentis new species, USNM 847234, spring on N side of Blanket Creek at Three Forks, Apache County.

Arizona: a, centrals (bar = 12.0 um); b, laterals and inner marginals (bar = 12.0 um); c, laterals and inner marginals (bar =

13.6 um); d, outer marginals ( bar = 7.5 um). FIGURE 29.-Penes of P. confluentis new species: a-d, USNM

847234. spring on N side of Blanket Creek at Three Forks, Apache

County, Arizona; e. USNM 847229, spring at Three Forks, Apache

County, Arizona.

FIGURE 30.-P. sancarlosensis new species and P. cochisi new species: a-g. P. sancarlosensis (a. USNM 859051, holotype. /1 springs W of Tom Niece Springs, Graham Colunty, Arizona; b. c. same locality, USNM 847226; d, e, USNM 847224, l arge spring N of

Bylas. Graham County, Arizona: f. g. USNM 847217, small spring near Bylas, Graham County, Arizona); h-k, P. cochisi (h, holotype, USNM 859055. springs at San Bernardino Ranch. Cochise

County, Arizona; i-k. same locality. USNM 847218) (bar = 0.75 mm).

FIGURE 31.-Radula of P. sancarlosensis new species. USNM

847226, springs W of Tom Niece Springs, Graham County, Arizona: a, b, centrals (bars = 15.0, 12.0 um); c, l aterals and inner marginals (bar = 8.6 um); d, laterals (bar = 15.0 um); e, inner marginals (bar = 6.0 um); f, outer marginals (bar = 8.6 um).

FIGURE 32.-Penes of P. sancarlosensis new species: a, USNM

847217, small spring near Bylas, Graham County, Arizona; b-d.

USNM 847226, springs W of Tom Niece Springs, Graham County,

Arizona.

FIGURE 33.-Radula of P. cochisi new species, USNM 847218, springs at San Bernardino Ranch, Cochise County, Arizona: a, centrals (bar = 9.1 um); b, l aterals (bar = 5.0 um); c. laterals and inner marginals (bar = 6.7 um); d, outer marginals (bar = 6.0 um). FIGURE 34.-Penes of P. cochisi new species, USNM 847218,

springs at San Bernardino Ranch, Cochise County, Arizona.

FIGURE 35.-P. thompsoni new species: a, holotype, USNM 859057,

Peterson Ranch Springs, Santa Cruz County. Arizona; b. USNM

847238, same locality; c, d, USNM 847221, Cottonwood Springs,

Santa Cruz County, Arizona; e. f, USNM 847228, Canelo Hills

Cienega, Santa Cruz County, Arizona; g, h. USNM 847223, Monkey

Springs, Santa Cruz County, Arizona; i-k, USNM 847227. Sheehy

Spring. Santa Cruz County. Arizona; 1, USNM 847251, Ojo Caliente.

Sonora. Mexico) (bar = 0.75 mm).

FIGURE 36.-Radula of P. thompsoni new species: a, centrals,

USNM 847238. Peterson Ranch Springs, Santa Cruz County, Arizona

(bar = 12.0 um); b. centrals. USNM 847221. Cottonwood Springs.

Santa Cruz County, Arizona (bar = 12.0 um); c. centrals. USNM

847227, Sheehy Spring, Santa Cruz County, Arizona (bar = 10.0

um); d, laterals, USNM 847238 (bar = 8.6 um); e, f, l aterals and

inner marginals, USNM 847238 (bars equal 6.7, 7.5 um); g, inner

marginals. USNM 847238 (bar = 10.0 um); outer marginals. USNM

847238 (bars equal 6.0 um).

FIGURE 37.-Penes of P. thompsoni new species: a, b. USNM

847238, Peterson Ranch Springs, Santa Cruz County, Arizona; c, d,

USNM 847228, Canelo Hills Cienega, Santa Cruz County, Arizona; e,

USNM 847221, Cottonwood Springs, Santa Cruz County, Arizona; f,

USNM 847223, Monkey Spring, Santa Cruz County, Arizona; g, USNM

847227, Sheehy Spring, Santa Cruz County, Arizona: h, USNM

847251, Ojo Caliente, Sonora, Mexico.

FIGURE 38.-Tryonia SPP•: a-e, T. gilae new species (a,

holotype, USNM 859059, small spring near Bylas. Graham County,

8 Arizona; b, c, same locality, USNM 847257; d, e, USNM 847253.

Cold Springs, Graham County, Arizona); f-h, T. quitobaquitae new species (f, holotype, USNM 859061, Quitobaquito Springs, Pima

County, Arizona; g, h, same locality, USNM 847256) ( bar = 0.75 mm).

FIGURE 39.-Dorsal aspect (without shell) of relaxed T. gilae new species. USNM 847257, small spring near Bylas, Graham County,

Arizona: a, female; b, male (Abbreviations: Cgp = posterior end

&capsule gland; Ct = ctenidium; Em = brooded embryo; I n intestine; Op = operculum; Pc = pericardium; Pn = penis; Sts = style sac).

FIGURE 40.-SEM micrographs of cephalic tentacles and penis of

Tryonia spp: a-c, T. gilae new species. USNM 847257. small spring near Bylas, Graham County, Arizona (a, left tentacle, bar = 100 um; b, right tentacle, bar = 120 um; c, close-up of ciliation on tentacle, bar = 20.0 um); d-f, T. quitobaquitae new species (d, left tentacle, bar = 75 um; e, right tentacle, bar = 75 um; f, distal half of penis, showing dense ciliation, bar = 50 um).

FIGURE 41.-Pallial cavity (viewed from (5e— right) of T. gilae new species, USNM 847257, small spring near Bylas, Graham County,

Arizona: Most of 6:§Lcapsule gland as been removed (position indicated by dashed lines) to reveal ( ctenidium and spermathecal duct (Abbreviations: Cg = capsule gland; Ct = ctenidium: I n = intestine; Mc = mantle collar; Os = osphradium;

Sd = spermathecal duct).

FIGURE 42.-Radula of Tryonia spp.: a-c, T. quitobaauitae new species, USNM 847256, Quitobaquito Springs, Pima County, Arizona

9 (a, centrals, bar = 8.6 um; b, l aterals and inner marginals, bar

= 6.7 um; c, outer marginals, bar = 5.0 um); d-i, T. gilae new species (a, centrals, USNM 847257, small spring near Bylas,

Graham County, Arizona. bar = 10.0 um; b. centrals. USNM 847253,

Cold Springs, Graham County, Arizona, bar = 7.5 um; c, l aterals,

USNM 847257. bar = 6.0 um; d. laterals. USNM 847253, bar = 5.0

UM; e, inner marginals, USNM 847253, bar = 12.0 um; f, outer marginals. USNM 847257. bar = 7.5 um).

FIGURE 43.-Anatomy of T. gilae new species, USNM 847257, small spring near Bylas, Graham County. Arizona: a. right aspect of prostate gland and adjacent organs and structures; b, right aspect of stomach (Abbreviations: Ast = anterior stomach chamber;

Au = auricle; Cc = caecal chamber; Cm = columellar muscle; Dgo = opening of digestive gland into stomach; I n = intestine; Ki kidney; Oes = oesophagus; Pr = prostate gland; Pst = posterior stomach chamber; Sts = style sac; Ve = ventricle).

FIGURE 44. SEM micrographs of penis of T. gilae new species, USNM 847257, small spring near Bylas. Graham County,

Arizona: a, dorsal aspect pf penis (bar = 200 um); b, close-up of distal end of above (bar = 75 um); c, d, dorsal aspect of penis base (bars equal 86, 100 um).

FIGURE 45.-Penes of T. gilae new species (a), USNM 847257, small spring near Bylas, Graham County, Arizona, and T. quitobaquitae new species (b), Quitobaquito Springs, Pima County,

Arizona: course ofof vas deferens within is indicated by dotted lines (Abbreviations: Ics = swelling on inner curvature; PI = penial l obe).

FIGURE 46.-The bursa copulatrix complex (viewed fromg right)

10 of T. puitobaquitae new species (a), USNM 847256, Quitobaquito

Springs, Pima County, Arizona, and T. gilae new species (b), USNM

847257, small spring near Bylas, Graham County, Arizona: Much of

pallial oviduct as bee cut away (position indicated by

dashed lines) to reveal underlying structures (Abbreviations: Ag

= albumen gland; Bu = bursa copulatrix; Cg = capsule gland; Dsr =

duct of seminal receptacle; In = intestine; Oes = oesophagus; Oov

opening of oviduct into albumen gland; Osd = opening of

spermathecal duct; Osr = opening of seminal receptacle duct into

spermathecal duct; Dv = oviduct; Sd = spermathecal duct; Sr

seminal receptacle; Sts = style sac).

FIG. 47.-Plot of scores on agi first two discriminant functions

derived from Pyrgulopsis shell data: a. malP i d.a.4.a—s-eZ- b. female:

.;part---...enk.t..L percent variation explained by each function is

indicated on respective axes; ellipses encircle positions of

topotypical specimens for each species (numbered as in Table 1);

1=P. deserta; 2=P. bacchus; 3=P. conicus; 4=P. morrisoni; 5= F.

montezumensis; 6=P. solus; 7=P. simplex; 8=P. glandulosus; 9=P.

confluentis; 10=P. sancarlosensis; 11=P. thompsoni; 12=P. — cochisi. Positions of ungrouped specimens (from additional -7 ----- populations) for three species are indicated by symbols.

FIG. 48.-Plot of scores on 1:!E! first two discriminant functions

derived from Pyrgulopsis anatomical data: a, male data set; b,

female data set: percent variation explained by each function is

indicated on respective axes; ellipses encircle positions of

topotypical specimens for each species (numbered as in Table 1);

1=P. deserta; 2=F. bacchus; 3=P. conicus; 4=P. morrisoni; 5= P.

11 montezumensis; 6=P. solus; 7=P. simplex; 8=P. glandulosus; 9=F. confluentis; 10=F. sancarlosensis; 11=P. thompsoni; 12=P. cochisi. Positions of ungrouped specimens (from additional populations) for three species are indicated by symbols.

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