Arizona Hydrobiidae
(Prosobranchia: Rissoacea)
*
ROBERT HERSHLER
and
J. JERRY LAND YE
m
i
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 459
SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION
Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first
Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a
program that included the following statement: "It is proposed to publish a series of reports,
giving an account of the new discoveries in science, and of the changes made from year to year
in all branches of knowledge." This theme of basic research has been adhered to through the
years by thousands of titles issued in series publications under the Smithsonian imprint,
commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the
following active series:
Smithsonian Contributions to Anthropology
Smithsonian Contributions to Astrophysics
Smithsonian Contributions to Botany
Smithsonian Contributions to the Earth Sciences
Smithsonian Contributions to the Marine Sciences
Smithsonian Contributions to Paleobiology
Smithsonian Contributions to Zoology
Smithsonian Folklife Studies
Smithsonian Studies in Air and Space
Smithsonian Studies in History and Technology
In these series, the Institution publishes small papers and full-scale monographs that report
the research and collections of its various museums and bureaux or of professional colleagues
in the world of science and scholarship. The publications are distributed by mailing lists to
libraries, universities, and similar institutions throughout the world.
Papers or monographs submitted for series publication are received by the Smithsonian
Institution Press, subject to its own review for format and style, only through departments of the
various Smithsonian museums or bureaux, where the manuscripts are given substantive review.
Press requirements for manuscript and art preparation are outlined on the inside back cover.
Robert McC. Adams
Secretary
Smithsonian Institution
S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 4 5 9
Arizona Hydrobiidae
(Prosobranchia: Rissoacea)
Robert Her shier
and
J. Jerry Landye
SMITHSONIAN INSTITUTION PRESS
Washington, D.C.
1988
ABSTRACT
Hershler, Robert, and J. Jerry Landye. Arizona Hydrobiidae (Prosobranchia: Rissoacea).
Smithsonian Contributions to Zoology, number 459, 63 pages, 49 figures, 6 tables, 1988.?A
diverse fauna of Arizona Hydrobiidae is documented as a result of recent collecting from
numerous springs in the state. The fauna is composed of 14 species in two genera, Pyrgulopsis
Call and Pilsbry and Tryonia Stimpson. Thirteen species are described as new herein and the
fourteenth represents a new state record. All have relatively restricted distributions, and two
are single-spring endemics. All congeners are allopatric to one another.
Stepwise canonical discriminant function analyses using sets of shell and anatomical data
confirmed the distinctiveness of the 12 species of Arizona Pyrgulopsis, as classification of
(grouped) topotypes was 89%-93% using shell data, and 100% using anatomical data.
Discriminant analyses also confirmed that differentiation among congeners largely involves
male anatomy (especially penial features), as this data set yielded the best separation of species
in plots of discriminant function scores.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's
annual report, Smithsonian Year. SERES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Hershler, Robert
Arizona Hydrobiidae (Prosobranchia: Rissoacea)
(Smithsonian Contributions to Zoology ; no. 4S9)
Bibliography: p.
Supc of Docs, no.: SI 127:459
1. Pyrgulopsis?Arizona?Classification. 2. Tryonia?Arizona?-Classification. 3. Mollusks?Classification.
4. Moltusks?Arizona?Classification. I. Landye, J. Jerry, n. Title. Ill Series
QL1.S54 no. 459 591s 87-600195 [QL43OO.5.H9] [594'32]
Contents
Page
Introduction 1
List of Recognized Species 1
Materials and Methods 3
Acknowledgments 6
Systematics 6
Key to Arizona Hydrobiidae 6
Family HYDROBIIDAE 7
Genus Pyrgulopsis Call and Pilsbry, 1886 7
Pyrgulopsis glandulosus, new species 8
Pyrgulopsis deserta (Pilsbry) 17
Pyrgulopsis bacchus, new species 21
Pyrgulopsis conicus, new species 21
Pyrgulopsis morrisoni, new species 21
Pyrgulopsis montezumensis, new species 23
Pyrgulopsis solus, new species 30
Pyrgulopsis simplex, new species 32
Pyrgulopsis confluentis, new species 32
Pyrgulopsis sancarlosensis, new species 35
Pyrgulopsis cochisi, new species 41
Pyrgulopsis thompsoni, new species 41
Genus Tryonia Stimpson, 1865 43
Tryonia gilae, new species 43
Tryonia quitobaquitae, new species 50
Morphometrics 50
Discussion 54
Addendum 58
Appendices 59
Literature Cited 63

Arizona Hydrobiidae
(Prosobranchia: Rissoacea)
Robert Her shier
and J. Jerry Landye
Introduction
Prosobranch snails of the family Hydrobiidae comprise a
diverse component of the inland aquatic fauna of the American
Southwest, with numerous locally endemic taxa found in
isolated springs, spring complexes, or drainage systems
(Taylor, 1966; Landye, 1973). In part due to difficulties of
collecting and studying these minute snails, they remain poorly
known. There are no published, modern, comprehensive,
systematic treatments of these faunas (but see Taylor, 1983),
and what little work has been done largely consists of isolated,
single-species descriptions based on studies of empty shell.
There is a clear and urgent need to address the systematics
of these faunas given the fact that many species are threatened
or endangered as a result of deterioration or disappearance of
natural aquatic habitats (Taylor, 1970a). As an illustration, 12
of 21 southwestern hydrobiid species currently being consid-
ered for federal listing as threatened or endangered (USDI,
1984) are undescribed.
To further this end, we present a systematic description of
Arizona Hydrobiidae. To date, not a single hydrobiid species
has been described from the state. Prior to 1969, the sole
published record of live collected hydrobiids in Arizona was
by Pilsbry and Ferriss (1909), who obtained Paludestrina
stearnsiana(l) from a single spring in the Huachuca Mountains.
In their monograph on Arizona mollusks, Bequaert and Miller
(1973) listed six additional such records for the family,
representing several possible species. Additional published
records of collections of Recent or subfossil shell are listed
by Taylor (1970b, 1975).
In the past 18 years the junior author has intensively
surveyed permanent Arizona waters and found a large
hydrobiid fauna, with estimated specific diversity ranging
Robert Hershler. Department of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560. J. Jerry Landye, Arizona
Game and Fish Department. HC 66. Box 2205. CornvilU. AZ 86325.
upwards to 18 species (see Landye, 1973, 1981; Williams et
al., 1985). The bulk of this collection, large alcohol holdings
from 38 localities in the state and peripheral areas (Figure 1),
was made available to the senior author, whose taxonomic
work is presented herein. Fourteen species are recognized,
representing two genera, Tryonia Stimpson, 1865, and
Pyrgulopsis Call and Pilsbry, 1886. Thirteen of the species are
new, and the fourteenth, Pyrgulopsis deserta (Pilsbry),
represents a new state record. Stepwise discriminant function
analyses using separate shell and anatomical data sets were
performed in order to examine patterns of variation and test
whether the 12 new, allopatric Pyrgulopsis species are distinct
from one another.
Only brief habitat description is provided, as a separate paper
treating ecological (and biogeographical) aspects of the fauna
will be published elsewhere. Hydrobiid localities in the state
are spring fed, and include spring pools, stream outflows, and
marshy wetlands known as cienegas (Hendrickson and
Minckley, 1985). Springs range from small seeps to Monte-
zuma Well, a limnocrene with maximum length of 112 m and
depth of 17 m (Cole and Barry, 1973). Snails are usually
restricted to headsprings and upper sections of outflows.
Pyrgulopsis typically occurs on rock or aquatic macrophytes
in moderate current, whereas Tryonia is usually restricted to
soft sediment in still water.
LIST OF RECOGNIZED SPECIES
Pyrgulopsis glandulosus Hershler, new species
Pyrgulopsis deserta (Pilsbry)
Pyrgulopsis bacchus Hershler, new species
Pyrgulopsis conicus Hershler, new species
Pyrgulopsis morrisoni Hershler, new species
Pyrgulopsis montezumensis Hershler, new species
Pyrgulopsis solus Hershler, new species
Pyrgulopsis simplex Hershler, new species
Pyrgulopsis sancarlosensis Hershler, new species
Pyrgulopsis confluentis Hershler, new species
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
i i i i i i i i i i i i i i i Kilometers
FIGURE 1.?Map of Arizona drainages, showing the 38 collecting sites numbered as in detailed list of collection
localities on facing page.
NUMBER 459
Collection Localities
Locality data includes appropriate information from USGS topographic quadrangle sheets
(7.5/15 minute series). Numbers in parentheses refer to localities in Figure 1; name of
quadrangle map follows each reference number.
United States
UTAH.?Washington County: Spring on Old Bastion Ranch (1), Hurricane
quadrangle, 0.0 km S, 0.05 km E from NE corner Sec. 13, T 42S, R. 15W;
Sam Adair Spring (2), St. George quadrangle, 0.8 km S, 0.6 km E from NW
corner Sec. 14, T. 42S, R. 15W; small spring, 70 m N of Interstate 15 (3), St.
George quadrangle, 0.1 km N, 0.1 km E from SW corner Sec. 15, T. 42S, R.
15W; Green Spring (4), St. George quadrangle, 0.6 km N, 0.7 km W from SE
corner Sec. 15, T. 42S, R. 15W; spring in Middleton (5), St. George quadrangle,
0.2 km W from NE corner Sec. 21, T. 42S, R. 15W; spring on west side of
Middleton (6), St. George quadrangle, 0.1 km S, 0.5 km E from NW corner
Sec. 21, T. 42S, R. 15W; spring on Red Hill (7), St. George quadrangle, 0.7
km N, 0.6 km E from SW corner Sec. 19, T. 42S, R. 15W.
ARIZONA.?Mohave County: Spring about 7.0 km E of Littlefield (8),
Litilefield quadrangle, 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
quadrangle, 0.5 km S, 0.2 km E from NW corner Sec. 3, T. 40N, R. 15W;
spring NE of Littlefield (10), Littlefield quadrangle, 0.8 km S, 0.5 km W from
NE corner Sec. 4, T. 40N, R. 15E; Grapevine Spring (11), Gypsum Hills
quadrangle, 0.0 km S, 0.4 km W from NE corner Sec. 26, T. 34N, R. 16W;
Burns Spring (12), Burns Spring quadrangle, 0.1 km S, 0.2 km W from NE
corner Sec. 9, T. 22N, R. 20W; Dripping Springs (13), Mount Nutt quadrangle,
0.6 km S, 0.7 km W from NE corner Sec. 4, T. 19N, R. 19W; Cool Spring
(14), Mount Nutt quadrangle, 0.5 km N, 0.5 km W from SE comer Sec. 20,
T. 19N, R. 19W.
Yavapai County: Tavasci Springs (15), Qarkdale quadrangle, 0.4 km N, 0.4
km E from SW corner Sec. 15, T. 16N, R. 3E; Page Springs (16), Page Springs
quadrangle, 0.1 km N, 0.3 km W from SE corner Sec. 23, T. 16N, R. 4E,
Montezuma Well (17), Lake Montezuma quadrangle, 0.9 km S, 1.1 km E from
NW corner Sec. 31, T. 15N, R. 6E; Brown Spring (18), Homer Mountain
quadrangle, 0.3 km N, 0.4 km W from SE comer of Sec. 23, T. 12N, R. 9E;
Fossil Springs (19), Strawberry quadrangle, 0.4 km N, 0.2 km W from SE
corner Sec. 14.T 12N.R.7E; Nelson Place Spring (21), Tule Mesa quadrangle,
6.0 km S, 4.3 km E from NW comer of quadrangle. Sec. 16, T. UN, R. 5E;
spring 150 m E of Nelson Place Spring (22), Tule Mesa quadrangle, 6.1 km
S, 4.8 km E from NW comer of quadrangle. Sec. 15, T. 1 IN, R. 5E.
Gila County: Spring near Strawberry (20), Strawberry quadrangle, 0.3 km
N, 0.3 km E from SW comer Sec. 22, T. 12N, R. 7E.
Apache County: Spring on N side of Blanket Creek at Three Forks (23), Big
Lake quadrangle, 0.5 km N, 0.2 km W from SE comer Sec. 6, T. 5N, R. 29E;
spring at Three Forks (24), Big Lake quadrangle, 0.2 km N, 0.0 km W from
SE comer Sec. 6, T 5N, R. 29E.
Graham County: Small spring near Bylas (25), Bylas quadrangle, 0.2 km
N, 0.3 km E from SW comer Sec. 17, T. 3S, R. 22E; spring near Gila River
bridge NW of Bylas (26), Bylas quadrangle, 0.1 km N, 0.3 km E from SW
comer Sec. 17, T. 3S, R. 22E; large spring N of Bylas (27), Bylas quadrangle,
0.1 km N, 0.3 km E from SW comer Sec. 28, T. 3S, R. 22E; springs W of Tom
Niece Springs (28), Fort Thomas quadrangle, 0.8 km S, 0.0 km W from NE
comer Sec. 21, T. 4S, R. 23 E; seeps W of Tom Niece Springs (29), Fort
Thomas quadrangle, 0.6 km S, 0.4 km W from SE comer Sec. 21, T. 4S, R.
23E; Cold Springs (30), Fort Thomas quadrangle, 0.1 km N, 0.5 km W from
SE comer Sec. 8, T. 5S, R. 24E.
Pima County: Quitobaquito Springs (31), Quitobaquito Springs quadrangle,
6.8 km S, 1.8 km W from NE comer of quadrangle, T. 17S, R. 7W.
Santa Cruz County: Cottonwood Springs (32), Sonoita quadrangle, 0.6 km
S, 0.1 km W from NE comer Sec. 33, T. 20S, R. 16E; Monkey Springs (33),
Sonoita quadrangle, 0.5 km N, 0.4 km E from SW comer Sec. 3, T. 21S, R.
16E; Canelo Hills Cienega (34), O'Donnell Canyon quadrangle, 0.6 km N, 0.1
km W from SE comer Sec. 32, T. 21S, R. 18E; Sheehy Spring (35), Lochiel
quadrangle, NW comer Sec. 12, T. 24S, R. 17E; Peterson Ranch Springs (36),
Huachuca Peak quadrangle, 0.5 km N, 0.6 km W from SE comer Sec. 3, T.
23S, R. 19E.
Cochise County: Springs at San Bernardino Ranch (38), College Peaks
quadrangle, 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).
Pyrgulopsis cochisi Hershler, new species
Pyrgulopsis thompsoni Hershler, new species
Tryonia gilae Hershler, new species
Tryonia quitobaquitae Hershler, new species
MATERIALS AND METHODS
MATERIAL EXAMINED.?Material from 38 localities (Figure
1), totalling 48 lots, was collected by fine-meshed hand sieve,
relaxed with menthol crystals, fixed in dilute (4%), buffered
formalin and preserved in 70% ethanol. Most of these lots
comprise more than 100 specimens. All this material is housed
in the United States National Museum collection in the
National Museum of Natural History, Smithsonian Institution
(USNM catalog numbers are referred to below), and other
relevant material from that collection and that of the Academy
of Natural Sciences of Philadelphia (ANSP) also was
examined.
MORPHOLOGIC STUDY.?Shells and animals were studied and
measured at x50 using WILD M-5 and M-7 dissecting
microscopes equipped with ocular micrometers. Shells, oper-
culae, and radulae were photographed using a Hitachi S-570
scanning electron microscope (SEM). To obtain intact bodies
for dissection, shells were~ decalcified by soaking in concen-
trated Bouin's Solution, and the remaining pellicle was
removed. Animals were dissected in dilute Bouin's Solution.
To examine penial morphology and ciliation patterns on the
penis and cephalic tentacles, animals were dried using a Dcnton
DCP-1 Critical Point Drier and then photographed by SEM.
Continuous and (to a lesser extent) meristic data were
obtained from shell and anatomy of selected series of sexed
specimens of Pyrgulopsis (Table 1) for use in discriminant
function analyses. Separate series were used for shell (n = seven
to eleven) and anatomical (n = five) study.
Shells were cleaned with Clorox to remove surface deposits
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
and oriented in the standard apertural aspect after counting of
whorls. Necessary points or aspects of shell outline were
pinpointed or traced onto paper using a camera lucida.
Distances between points were then determined using a
fine-scaled millimeter ruler. The following characters were
measured (Figure 2a) or counted:
number of whorls (NW)
shell height (SH)
shell width (SW)
aperture height (AH)
aperture width (AW)
height of body whorl (LBW)
width of body whorl (WBW)
length of line connecting sutures at posterior ends of body
and penultimate whorls (AB)
length of perpendicular bisector of line segment AB,
extended to edge of penultimate whorl (CD)
The following ratios were formed using the above data:
shell height/shell width (SH/SW)
aperture height/shell height (AH/SH)
body whorl length/shell height (LBW/SH)
whorl convexity (CD/AB), following Kohn and Riggs (1975:352)
Body and gonad lengths were measured with the animal
pinned in an apical aspect, with the former representing length
from mantle collar to posterior tip of digestive gland. The penis
was cut from the neck and measured in dorsal aspect. The
pallial roof and visceral coil were removed from the head/foot
and separated from one another along the end of the pallial
cavity. Pallial structures were measured with the pallial roof
flattened and pinned. Penial measurements are shown in Figure
2b as an example of continuous anatomical data. The following
counts and measurements were made:
Both Sexes
body length (BL)
osphradium length (OSL)
osphradium width (OSW)
distance from posterior end of osphradium to posterior tip
of ctenidium (OCT)
ctenidium length (CTL)
ctenidium width (CTW)
number of ctenidial filaments (FI)
Females
ovary length (OL)
capsule gland length (CGL)
capsule gland width (CGW)
albumen gland length (AGL)
albumen gland width (AGW)
diameter (anterior-posterior) of anterior coil of oviduct (COL)
distance from anterior end of oviduct coil to posterior end
of capsule gland (COP)
maximum diameter of anterior portion of oviduct (COD)
bursa copulatrix length (BUL)
bursa copulatrix width (BUW)
width of bursa copulatrix duct (BDW)
distance from posterior end of albumen gland to posterior
tip of bursa (BAG)
TABLE 1. List of Pyrgulopsis populations receiving morphometric study of
shell and anatomy. USNM collection numbers are followed by locality numbers
(see Figure 1). Specimens from type-localities are indicated with asterisks.
(Plus = data taken, dash = no data taken.)
Species and locality
P. deserta
?847202 (9)
847206 (8)
P. bacchus
?847203 (1 )
P. conicus
?847237 (13)
847201 (14)
P. morrisoni
?847231 (16)
847220 (15)
P. montezumensis
?847233 (17, swallet)
847230 (17, outflow)
P. solus
?850290 (18)
P. glandulosus
?847205 (21)
P. simplex
?847236 (20)
847222 (19)
P. confluentis
?847234 (23)
847229 (24)
P. sancarlosensis
?847226 (28)
847217 (25)
847224 (27)
P. thompsoni
?847238 (36)
847221 (32)
847227 (34)
847228 (34)
P. cochisi
?847218 (38)
Shell Anatomy
+ +
+
+ +
+ +
+
+ +
+
+ +
+
+ +
+ +
+ +
+
+ +
+
+ +
+ +
+
+ +
+ +
+ +
+ +
+ +
seminal receptacle (body plus duct) length (SRL)
seminal receptacle width (SRW)
Males
testis length (TS)
prostate gland length (LPR)
prostate gland width (WPR)
length of penis from base to tip of penial lobe (L2)
penis width (at base) (PW)
distance from base of penis to base of penial filament (L3)
penial filament length (LFI)
penial filament width (WFI)
number of glandular ridges on penis (GLR)
The following ratios were formed using these anatomical
data:
Both Sexes
osphradium length/ctenidium length (OSL/CTL)
NUMBER 459
a
WBW
SH
LBW
L2
L3
-AW
SW- PW-
FIGURE 2.?Shell (a) and penial (b) measurements. Points: a, suture at posterior end of penultimate whorl; b,
suture at posterior end of body whorl; c, mid-point of line segment ab; d, intersection of perpendicular bisector
of ab and edge of whorl (Abbreviations: AH = aperture height; AW = aperture width; LBW = height of body
whorl; LFI - length of penial filament; L2 = penis length; L3 = length from proximal end of penis to distal edge
of filament base; PW - penis width; SH = shell height; SW = shell width; WFI = width of penial filament;
WBW = width of body whorl.)
distance from posterior tip of osphradium to ctenidium tip/ctenidium
length (OCT/CTL)
ctenidium length/body length (CTL/BL)
Females
ovary length/body length (OV/BL)
capsule gland length/albumen gland length (CGL/AGL)
oviduct coil diameter/albumen gland length (COL/AGL)
bursa length/albumen gland length (BUL/AGL)
distance from albumen gland tip to bursa tip/bursa length
(BAG/BUL)
seminal receptacle length/bursa length (SRL/BUL)
Males
tcstis lengih/body length (TS/BL)
prostate length/body length (LPR/BL)
distance from base of penis to base of penial filament/penis
length (L3/L2)
length of filament/penis length (LFI/L2)
filament length/width (WFI/LFI)
The total character breakdown is as follows: male and female
shell, nine raw characters, four ratios (totaling 13 characters);
male anatomy, 16 and eight (24); female anatomy, 21 and nine
(30).
DATA ANALYSIS.?As discussed by Mayr (1963) and Wiley
(1981), assessment of systematic status of differentiated
allopatric populations 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, 355-365) and was therefore used to
examine patterns of variation among the 12 allopatric
Pyrgulopsis species described below.
We employed canonical discriminant function analysis, also
known as canonical variates analysis when applied (as in this
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
case) to more than two groups, as this technique is particularly
suitable when one seeks to evaluate distinctiveness of a priori
designated taxa. Unlike principal component analysis, discri-
minant analysis provides statistical evaluation of specimen
classification. We stress that these analyses were for the
purposes of description and exploration of the data set and are
not intended as inferential.
MAHAL stepwise procedure was used in discriminant
analyses, in which variables are selected on the basis of
providing the largest Mahalanobis distance between the closest
pair of groups. Stepwise procedure was used, as it allows
recognition of the most effectively discriminating variables.
Topotypes of each species were considered as 12 separate
groups, with other populations input as ungrouped to prevent
biases resulting from having data available for varying numbers
(one to four) of populations for the various species. All
populations used 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 male and
female shell data, as sexual dimorphism in size is evident in
at least one population for 10 of 12 species and in 11 of 23
total populations studied (Appendix 1). The pooled within-
groups covariance matrix was used to compute probabilities
of group membership.
Means for all raw counts, measurements, and ratios except
one (OSW, females) were heterogeneous among the 12 sets
of topotypes. Ratios were used in the analyses as a way of
dealing with shape. Initial discriminant analyses using all
variables indicated that significant separation (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 variables. Selection of additional
variables no further separated groups, and eventually (after 16
to 20 variables added) resulted in decreased separation. To
avoid possible confusion of patterns generated by addition of
later variables, discriminant analyses were rerun using subsets,
with each consisting of all variables selected (in initial
analyses) to the point at which consistent significant separation
occurred between closest groups. The reduced variable sets
thus selected were as follows: male anatomy (11 variables),
CTW, TS/BL, L2, BL, L3/L2, GLR, LFI, WFI, CTL, LPR/BL,
L3; female anatomy (13), CTL, BAG, BUW, COP, CGW, FI,
AGL, OV, SRL/BUL, OSL, BDW, OCT, OSL/CTL. For shell
data (consisting of relatively few variables), decreased
separation of closest groups occurred after every step in initial
analyses, and therefore runs utilizing all variables were used.
Descriptive statistics were generated, and analyses of
variance (ANOVA), t-tests, and stepwise discriminant function
analyses were performed using the SPSS-X program, subpro-
grams BREAKDOWN, T-TEST, and DISCRIMINANT.
ACKNOWLEDGMENTS
We thank Drs. G.M. Davis, Academy of Natural Sciences
of Philadelphia (ANSP) and W. Pratt (Museum of Natural
History, University of Nevada at Las Vegas) for lending
material. Paul Greenhall (Department of Invertebrate Zoology,
National Museum of Natural History (NMNH), Smithsonian
Institution) measured shells and provided other diverse
technical assistance. Dr. Lee-Ann Hayek and Cindy Carmen
(Office of Information Resources Management, Smithsonian
Institution) ran appropriate computer programs and helped
greatly with interpretation of data analyses. Molly Kelly Ryan
(Staff Illustrator, Department of Invertebrate Zoology, NMNH,
Smithsonian Institution) prepared most of the final illustrations
and plates. The SEM micrographs were taken with the help of
Walter Brown, Heidi Wolf, and Susann Braden of the SEM
Laboratory, NMNH, Smithsonian Institution. Drs. S.M. Cham-
bers, G.M. Davis, W.L. Minckley, and W.F. Ponder provided
useful criticism of the manuscript.
Hershler's participation in the project was funded, in part,
by Arizona State University. Part of Landye's fieldwork was
supported from 1964 through 1971 by grants to Dr. W.L.
Minckley, and later by U.S. Fish and Wildlife Service
contracts.
Systematics
Accounts for each species consist of provision of common
name, synonymy, material examined, diagnosis, description,
remarks (where applicable), type-locality, distribution, and
etymology (where applicable). Designation of common names
largely follows that of Landye (1981) and Williams et al.
(1985). Most species were recognized in unpublished reports
by Landye (1973, 1981). Our synonomies include references
to these works only when identities of his taxa are unclear.
Congeners are not greatly divergent in morphology and
expanded descriptions are provided for only one species per
genus, with other descriptions emphasizing distinctive features.
Such features involve (in order of importance) penial
morphology, shell, head-foot and penial pigmentation, and
morphology of the pallial oviduct and bursa copulatrix
complex. Radular data are in Table 2. Shell and anatomical
data for Pyrgulopsis species are in Appendices 1-3.
Key to Arizona Hydrobiidae
1. Shell elongate-conic to turretcd; penis with glandular, papillae-like lobes (Tryonia).
2
Shell globose to elongate-conic; penis bilobed, with glandular ridges (Pyrgulopsis).
NUMBER 459 7
2. Adult shell height 1.9-3.3 mm; penis with two lobes on inner curvature near distal
tip and single, enlarged lobe on outer curvature at base
Tryonia gilae, new species
Adult shell height, 1.4-1.8 mm; penis with single lobe on inner curvature near distal
tip and enlarged lobe on inner curvature at base
Tryonia quitobaquitae, new species
3. Penial filament narrow, short (less than 30% of penis length); dorsal penial surface
with two or more glandular ridges 4
Penial filament wide, long (greater than 40% of penis length); dorsal penial surface
with one or no glandular ridges 6
4. Ventral penial surface without accessory crests
Pyrgulopsis montezumensis, new species
Ventral penial surface with accessory crest(s) 5
5. Dorsal penial surface with at least two elongate ridges; ventral surface with two
accessory crests Pyrgulopsis glandulosus, new species
Glandular ridges on dorsal penial surface circular; ventral surface with single
accessory crest Pyrgulopsis confluentis, new species
6. Penial lobe reduced or absent 7
Penial lobe prominent 8
7. Adult shell height, 1.3-1.7 mm; penial lobe absent; dorsal and ventral penial surfaces
each with single glandular ridge Pyrgulopsis cochisi, new species
Adult shell height, 1.7-3.2 mm; penial lobe reduced; ventral penial surface with
single glandular ridge Pyrgulopsis thompsoni, new species
8. Dorsal penial surface with glandular ridge(s) 9
Dorsal penial surface without glandular ridges 11
9. Dorsal surface of penial filament with elongate glandular ridge
Pyrgulopsis deserta (Pilsbry)
Dorsal surface of penial filament without elongate glandular ridge 10
10. Penis short (about 13%) relative to body length; penial filament 40%-50% of penis
length Pyrgulopsis solus, new species
Penis long (about 33%) relative to body length; penial filament greater than 67% of
penis length Pyrgulopsis sancarlosensis, new species
11. Penial lobe bi- or trifurcate; ventral penial surface with single elongate or several
circular glandular ridges Pyrgulopsis bacchus, new species
Penial lobe simple; ventral penial surface with single, circular glandular ridge . . 12
12. Penial filament length greater than 50% of penis length, typically extending beyond
distal tip of penial lobe Pyrgulopsis simplex, new species
Penial filament length 50% or less than penis length, rarely extending beyond distal
tip of penial lobe 13
13. Penis length about 15% of body length; penial filament length less than three times
filament width Pyrgulopsis conicus, new species
Penis length 24%-30% of body length; penial filament length about three and a half
times filament width Pyrgulopsis mornsoni, new species
Family HYDROBDDAE REMARKS.?A detailed generic description (with syno-
nymies) is provided by Hershler and Thompson (1987), in
Genus Pyrgulopsis Call and Pilsbry, 1886 which Marstonia F.C. Baker, 1926, Fontelicella Gregg and
Taylor, 1965, Microamnicola Gregg and Taylor, 1965,
DIAGNOSIS.?Shell globose to ovate-conic, 1.2 to 8.0 mm Natricola Gregg and Taylor, 1965, and Mexistiobia Hershler,
high. Aperture simple, umbilicus absent to open. Teleoconch 1985 are considered synonymous with Pyrgulopsis. The genus
smooth or unicarinate on periphery. Penis with one to fifteen occurs in much of eastern North America as well as in the
glandular ridges. Penial lobe small relative to size of penis, Southwest and northern Mexico and comprises 35 (including
filament narrow. the below) described species as well as an additional 20 to 25
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.?Generalized cusp formulae for the four tooth types.
Species
Pyrgulopsis glandulosus
P. deserta
P. bacchus
P. conicus
P. morrisoni
P. montezumensis
P. simplex
P. solus
P. confluentis
P. sancarlosensis
P. cochisi
P. thompsoni
Tryonia gilae
T. quitobaquilae
Central
5-1-5
1-1
4-1-4(5)
1-1
3(4,5)-l-3(4,5)
1-1
4-1^4
1-1
6<7>-l-6(5)
1-1
8-1-8
1-1
5-1-5
1-1
5(6)-l-5(6)
1-1
4-1-5
1-1
3(5)-l-3(5)
1-1
4-l^t
1-1
4(5,6>-l-5(6)
1-1
5(6>-l-5(6)
2-2
6-1-6
1-1
Lateral
3-1-3(4)
3(4>-l-3
2(3)-l-2
3(4)-l-3
3-1-3(4)
3(4,5)-l-5(6)
3-1-5
3(4,5)-l-4(5)
3-1-4
2(3)-l-3
3-1-3
3-1-3(4)
4(5,6)-l-5(6)
3-1-4
Inner
marginal
23
17-19
18-19
19
22-26
27-31
23-24
21
24-32
16-22
22-23
20-24
24-30
23
Outer
marginal
31
19-21
30-31
25-28
27-29
28-30
28-30
25-26
23
23-25
22-23
26-32
28-31
26
undescribed species from the Southwest. The closest relative
of the genus appears to be Cincinnatia Pilsbry, 1891, which
is restricted to eastern North America. Cincinnatia differs from
Pyrgulopsis in having a larger and broader shell, and a penis
with a much smaller penial filament (relative to size of penis)
and more glandular ridges.
Pyrgulopsis glandulosus Hershler, new species
Verde Rim springsnail
FIGURES 3a-c, 4-9, \0a,b
Fontelkella species.?Williams etal., 1985:19.
MATERIAL EXAMINED.?ARIZONA. Yavapai County: Nelson
Place Spring, USNM 859047 (holotype), USNM 859048 (5
paratypcs), USNM 847205, JJ. Landye and G. Edwards, 28
Sep 1973; spring 150 m E of Nelson Place Spring, USNM
847232, J J . Landye and G. Edwards, 28 Sep 1973.
DIAGNOSIS.?An ovate-shelled, large size (shell height, 2.0
to 2.8 mm) species. Penis elongate, massive (41% of body
length), filament short relative to penis length. Dorsal penial
surface with several glandular ridges, two or more of which
are elongate; ventral surface with additional ridges borne on
accessory crests. Holotype (Figure 3a) height, 2.80 mm; width,
1.92 mm; whorls 4.0.
DESCRIPTION.?Shell white-clear, transparent; periostracum
light brown, thin, covering much of shell surface, or absent.
Shell (Figure 3a-c) about a third taller than wide. Whorls, 3.5
to 4.0, convex and slightly shouldered, sutures slightly
impressed. Spire outline convex, largely due to allometric
translation of body whorl. Sexual dimorphism significant
(Appendix 2) with females larger than males. Protoconch
slightly protruding, with 1.25 partly pitted whorls. Body whorl
relatively inflated and large (about 83% of shell height).
Aperture ovate, moderate in size, occupying about half of shell
height and 83% of body whorl height, somewhat angled above,
rounded below, typically loosened from body whorl. Inner lip
slightly thickened, moderately reflected, and near straight.
Outer lip thin and broadly convex. Apertural plane nearly
parallel to coiling axis with long axis tilted 20? to 30? relative
to coiling axis. Umbilicus moderately to broadly open.
Teleoconch sculpture consisting of strong growth lines. Amber,
horny, paucispiral operculum about one and a half times as
long as wide and with three whorls.
Snout (Figures 4/ 5c, la) longer than wide, fairly thickened,
and terminating distally with fleshy lips. Cephalic tentacles
(Tn) narrow, slightly less than twice as long as snout,
somewhat expanded at tips. Tentacle surface with irregular
NUMBER 459
FIGURE 3.?Pyrgulopsis glandulosus, new species: a, holotype, USNM
859037, Nelson Place Spring, Yavapai County, Arizona; b,c, same locality,
USNM 847205. Pyrgulopsis confluentis, new species: 4 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 4.?Right lateral aspect of relaxed P. glandulosus, new species (without
shell), USNM 847205, Nelson Place Spring, Yavapai County, Arizona: darkly
pigmented epithelia appear black, and glandular ridges in penis screened.
(Abbreviations: Ft = foot, Grn = white granules; He = head, Opl =
operculigerous lobe, Pn = penis; Sep = subepithelial pigment; Tn = cephalic
tentacle.)
10 SMITHSOXIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 5.?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 urn; b, dorsal aspect
of penis, bar = 0.33 mm; c, right tentacle, showing sparse ciliation, bar = 176 \un; d, left tentacle, bar = 136 \xm.
N'UMBER 459 11
tufls of cilia (Figure 5b,c), sometimes forming a short tract on
dorsal surface of left tentacle. Slightly swollen outer edges of
tentacle bases containing black eyes. Elongate foot thickened,
posteriorly tapered. Fairly straight anterior edge of foot with
central pore through which twelve to fifteen elongate pedal
glands discharge.
Epithelial melanic pigment usually absent from head/foot
although occasional snails may have light to fairly dark brown
patches covering most of snout (posterior to distal lips). Dorsal
surface of visceral coil covered with melanic pigment (Figure
4), although stomach region typically less pigmented or even
lacking pigment. Light grey to black subepithelial pigment
sometimes present as small patches just distal to bases of
tentacles and on "neck" behind tentacles, as broad triangular
patch on sides of head/foot (Figure 4, Sep), and as irregular
patch on operculigerous lobe. Similar pigment present on right
side of mantle collar. Opaque white granules densely clustered
just behind eyes (Figure 4, Grn), with small clusters sometimes
occurring along proximal half of tentacles as well. Dark,
brown-black subepithelial pigment concentrated in proximal
portion of penial filament, extending somewhat to rest of penis
(Figure 4). Ventral pigmentation visible dorsally as in Figure 4.
Pallial cavity (Figure 6a,b) with large ctenidium (Ct) with
16 to 20 narrow filaments extending along most of pallial
cavity (and 24% to 28% of body length), and extending
anteriorly to mantle collar (Me). Widest filaments centrally
positioned and brushing against intestine (in). Osphradium
(Os) fairly elongate, filling 22% to 33% of ctenidium length,
positioned slightly posterior to middle of ctenidium. Kidney
(Ki) with small opening (Oki) fringed by obvious lips.
Buccal mass short, filling only anterior half of snouL Paired,
tubular, salivary glands dorsal to nerve ring. Oesophagus
externally simple, and narrow. Pair of chitinous jaws
positioned at anterior end of buccal cavity.
Radula (Figure 7) typically taenioglossate, with trapezoidal
central teeth, each having single pair of basal cusps. Lateral
angles of central tooth well developed, projecting about 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 former
having only few more cusps than latter (Table 2). Stomach
(Figure 6c, Ast, Pst) and style sac (Sts) about equal in length,
former with small caeca! chamber (Cc) along its posterior edge.
Posterior stomach chamber with single opening to digestive
gland (Dgo) (posterior to oesophageal opening). Digestive
gland covering posterior half of stomach, and extending
(ventral to gonad) almost to posterior tip of visceral coil.
Intestine (In) with slight bend in pallial cavity (Figure 6b).
Testis (Figure 6d, Ts) consisting of simple lobes, with
anterior end lying against posterior edge of stomach, and filling
about three-quarters of digestive gland length and 31 % of body
length. Seminal vesicle (Figure 6d, Sv) exiting near anterior
end of testis and consisting of a few thickened coils ventral to
testis and partly covering dorsal stomach surface. Prostate
gland (Figure 6e, Pr) simple, small (16%) relative to body
length, largely posterior to pallial cavity. Anterior vas deferens
(Figure 6e, Vd) exiting from mid-line of anterior end of
prostate gland and extending along columellar (right) side of
pallial cavity roof before entering "neck" posterior to penial
attachment area (Figure 6a).
Penis (Figures 4, 5a,b, 6a, 8, 9) protruding well anterior to
mantle collar (Figure 5a), flattened, broadly attached to neck
well posterior to snout (Figure 6) and extending anteriorly from
attachment area without coiling (Figure 6a, Pn). Penis much
longer than wide, often expanding in width distal to base.
Penial filament (Figure 8, Pf) one-quarter of penis length,
positioned at about two-thirds of penis length, narrow
(length/width, 25%), and rarely projecting distal to penial lobe
(Figure 8, Plo), which tapers and sometimes has distinctive
terminal process (Figure $b,d). Vas deferens (Vd) extending
as fairly straight tube along edge of penis (Figure 8a), and
opening at tip of penial filament.
Penial surface with four to seven slightly elevated, glandular
ridges, consisting of paired rows of small glands, each
discharging through central slit (Hershler, 1985, fig. 8b,c).
Dorsal penial surface with two elongate ridges, one positioned
largely along edge containing penial filament, and other located
along opposite edge, sometimes curving toward filament
(Figures 8, 9). One to three additional and smaller ridges on
same surface, typically positioned on tip of penial lobe (visible
from ventral aspect also) and between two elongate ridges.
Ventral penial surface with one to three ridges located on two
accessory crests. Crest located closest to distal edge of penis
somewhat swollen and bulging from penial surface, whereas
other crests typically appearing as folds flattened against penis.
Ovary (not figured) simply lobular, abutting anteriorly
against posterior edge of stomach, and filling 17% of body
length. Oviduct passing over stomach surface before disappear-
ing under (to left of) pallial oviduct. Left lateral aspect of pallial
oviduct and bursa copulatrix complex shown in Figure \0a,b.
Albumen gland (Ag) highly glandular and somewhat (104%)
longer than capsule gland (Cg), which has posterior, yellow
section and (smaller) anterior, clear section. Capsule gland
opening (Cga) subterminal and fairly long and wide. Anterior
to opening, presumed pathway of egg capsules fringed by thin,
vestibule-like extension of glandular tissue from capsule gland
(Figure 10a, Vsb), as typical of genus. Oviduct (Ov) 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, positioned on left surface
of bursa, often narrow and indistinguishable from its duct;
length of body plus duct almost equal (83%) to bursa length.
Bursa (Figure \0b) moderately large (43% of albumen gland
length), pear-shaped, and lying against left side (near ventral
Emc
i 1 0.5 mm
FIGURE 6.?Anatomy of P. glandulosus, new species, USNM 847205, Nelson Place Spring, Yavapai County,
Arizona: a, head and contents of pallial cavity, which were exposed by slitting pallial roof mid-dorsally; b,
ctenidium and adjacent structures; c, right lateral aspect of stomach; d, coiled seminal vesicle (testis removed);
e, right lateral aspect of the prostate gland. (Abbreviations: Ast = anterior stomach chamber, Cc = caecal
chamber. Cm = columellar muscle; Ct = ctenidium; Dgo - opening of the digestive gland into the stomach;
Emc = posterior end of pallial cavity; In = intestine; Ki = kidney; Me - mantle collar, Oes = oesophagus;
Oki = kidney opening; Os = osphradium; Pn = penis; Pr = prostate; Pst = posterior stomach chamber;
Sn = snout; Sts = style sac; Sv = seminal vesicle; Tn = cephalic tentacle; Ts = testis; Vd = anterior vas deferens;
Vdl = posterior vas deferens.)
NUMBER 459 13
FIGURE 7.?Radula of P. glandulosus, new species, USNM 847205, Nelson Place Spring, Yavapai County,
Arizona: a, centrals, bar = 13.6 Jim; b, laterals, bar = 15.0 |im; c, laterals and inner marginals, bar = 15.0 (im;
d, outer marginals, bar = 5.0 fim.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Plo
FIGURE 8.?Penial variation of P. glandulosus, new species, USNM 847205, Nelson Place Spring, Yavapai
County, Arizona: Dorsal aspects to left, ventral aspects to right; screened areas indicate glandular ridges.
(Abbreviations: Ace = accessory crest; Pf = penial filament; Plo = penial lobe; Vd = vas deferens.)
NUMBER 459 15
FIGURE 9.?Penial variation of P. glandulosus, new species: a,b, USNM 847205, Nelson Place Spring, Yavapai
County, Arizona; c, USNM 847232, spring 150 m E of Nelson Place Spring, Yavapai County, Arizona. (Left =
dorsal aspect; right = ventral aspect; screened areas - glandular ridges.)
edge) of albumen gland. Two-thirds of bursa length posterior
to gland. Thin, narrow duct of bursa tightly pressed to albumen
gland.
REMARKS.?Pyrgulopsis glandulosus groups with a number
of Arizona and New Mexico species (the latter undescribed,
see Taylor, 1983) having large, ovate shells and penes with
small filament and relatively numerous glandular ridges. On
the basis of penial morphology, the species is most similar to
P. montezumensis, which also occurs in Verde River drainage,
but differs in having a larger penis and accessory crests on
Emc
Dbu
Cg
FIGURE 10.?Pallia] oviducts of Pyrgulopsis species, 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 pallial cavity, with mantle edge indicated by dashed 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; e, P. conicus, new species, USNM 847237, Dripping Springs, Mohave County, Arizona;/ P. morrisoni,
new species, USNM 847231, Page Springs, Yavapai County, Arizona; g, P. montezumensis, new species, USNM
847233, Montezuma Well, Yavapai County, Arizona. (Abbreviations: Ag = albumen gland; Bu = bursa
copulatrix; Cg = capsule gland; Cga = capsule gland opening; Cm = columellar muscle; Dbu = duct of bursa;
Emc - posterior end of pallial cavity; In = intestine; Vsb = capsule gland vestibule.)
NUMBER 459 17
FIGURE 11.?P. deserta (Pilsbry): 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. P.
bacchus, new species: e, USNM 859037, holotype. Grapevine Spring, Mohave County, Arizona; fg, USNM
847203, same locality. (Bar = 0.75 mm.)
ventral penial surface.
TYPE-LOCALITY.?Nelson Place Spring, Yavapai County,
Arizona.
DISTRIBUTION.?Nelson Place Spring complex, consisting
of two springs separated by 150 m that form the headwaters
of Sycamore Creek.
ETYMOLOGY.?From Latin glandis, referring to the highly
glandular nature of the penis in this species.
Pyrgulopsis deserta (Pilsbry)
Virgin springsnail
FIGURES 10C, lla-d, I2a-c, 13
Amnicola deserta Pilsbry, 1916:111-112, figs. 8, 9.
New genus, new species.?Landye, 1973:48.
"Amnicola" deserta.?Landye, 1980:70.
Undescribed species.?Landye, 1980:70.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 12.?Radula of P. deserta (Pilsbry), USNM 847202, spring upstream on Virgin River from Littlefield,
Mohave County, Arizona: a, centrals, bar = 12.0 \m; b, laterals and inner marginals, bar = 15.0 ^m; c, outer
marginals, bar = 17.6 Jim. P. bacchus, new species, USNM 847203, Grapevine Spring, Mohave County, Arizona:
d, centrals, bar = 20.0 Jim; e, centrals, laterals, inner marginals, bar = 38 jam;/ laterals, inner and outer marginals,
bar = 38 urn
MATERIAL EXAMINED.?UTAH. Washington County: spring
on Old Bastion Ranch, USNM 847250, JJ. Landye, 21 Feb
1977; small spring, 70 m N of Interstate 15, USNM 847247,
JJ . Landye, 21 Feb 1977; Green Spring, USNM 847242, JJ .
Landye, 21 Feb 1977; spring in Middleton, USNM 847210,
JJ . Landye, 21 Feb 1977; spring on Red Hill, USNM 847239,
JJ . Landye, 30 Aug 1973; spring on west side of Middleton,
USNM 847244, J J . Landye, 20 Feb 1977; Sam Adair Spring,
USNM 847245, JJ . Landye, 21 Feb 1977; ANSP 12112
(holotype). ARIZONA. Mohave County: spring just E of
Littlefield, USNM 847206, JJ . Landye, 23 Feb 1977; spring
upstream on Virgin River from Littlefield, USNM 847202, JJ.
Landye, 31 Aug 1973; spring NW of Littlefield, W. Rinne, 17
Dec 1970.
DIAGNOSIS.?A small species (shell height, 1.2 to 2.0 mm)
with globose to ovate-conic shell. Penis moderate size, with
enlarged filament bearing (on its dorsal surface) elongate
glandular ridge; ventral penial surface with two to three
additional ridges. Holotype (Figure l la) height, 2.16 mm;
width, 1.57 mm; whorls, 4.0.
DESCRIPTION.?Shell with 3.5 to 4.25 whorls, varying in
shape from globose, with well-rounded, unshouldered whorls
and relatively large (60% of shell height) aperture (Figure
\\c,d) to ovate-conic, with fairly flattened whorls and smaller
NUMBER 459 19
FIGURE 13.?Penial variations of P. deserta (Pilsbry): 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. (Left = dorsal aspect; right =
ventral aspect; screened areas = glandular ridges.)
aperture (Figure \\a,b). Shape variation evident not only
among, but also within populations, with enlarged conical
specimens sometimes forming a distinct size class. Sexual
dimorphism significant (males larger than females) in one of
two populations studied. Aperture often partly or completely
loosened from body whorl. Umbilicus either absent or
chink-like to entirely open.
Melanic pigmentation of head-foot varying from dark
throughout (except for tentacle and snout tips, and sole of foot)
in some populations, to a light dusting on all surfaces or even
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 14.?Protoconchs and opercula of Arizona hydrobiids: a, protoconch, P. montezumensis, new species,
USNM 847233, Montezuma Well, Yavapai County, Arizona, bar = 136 urn; b, proloconch, P. bacchus, new
species, USNM 847203, Grapevine Spring, Mohave County, Arizona, bar = 136 fim; c, protoconch, Tryonia
gilae, new species, USNM 847257, small spring near Bylas, Graham County, Arizona, bar = 75 Jim; d,
operculum, P. montezumensis, same lot as above, bar = 0.33 mm; e. operculum, P. sancarlosensis, new species,
USNM 847226, springs W of Tom Niece Springs, Graham County, Arizona, bar = 0.27 mm; / operculum, T.
gilae, same lot as above, bar = 176 urn.
absent in others. Penial filament pigmentation ranging from
dark throughout to near-absent.
Ctenidial filaments few, seven to 10. Radula (Figure 12)
characterized by relatively few cusps on central and marginal
teeth. Testis relatively small (31% of body length); prostate
gland relatively large (20% of body length). Penis elongate
(Figure 13), filament 74% of penis length, extending beyond
tip of penial lobe, and fairly thickened (length/width, 42%).
Lobe relatively small, sometimes tapering. Glandular ridges,
three to four. Ridges on ventral surface positioned on or just
behind penial lobe. Bursa (Figure 10c) enlarged (84% of
albumen gland length). Seminal receptacle small (40%) relative
to bursa length.
REMARKS.?This snail does not closely resemble any other
Arizona form, but is similar to an undescribed species also from
Virgin River drainage, southwestern Utah.
TYPE-LOCALITY.?Pilsbry (1916:111) 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
holotype.
DISTRIBUTION.?Springs along Virgin River in southwestern
Utah and northwestern Arizona.
NUMBER 459 21
Pyrgulopsis bacchus Hershler, new species
Grand Wash springsnail
FIGURES 104 Ue-g, \2d-f, 146, 15
Fontelicella species.?Williams et al., 1985:32.
MATERIAL EXAMINED.?ARIZONA. Mohave County: Grape-
vine Spring, USNM 859037 (holotype), 859038 (4 paratypes),
847203, J J . Landye, 4 Jun 1980.
DIAGNOSIS.?A large size (shell height, 2.3 to 3.1 mm),
ovate-shelled species. Penis small (13% of body length), with
thickened, unpigmented filament; penial lobe enlarged and
often bifurcate. One to three glandular ridges on ventral penial
surface, along distal edge of penial lobe. Holotype (Figure
lie) height, 2.41 mm; width, 1.78 mm; whorls, 4.0.
DESCRIPTION.?Shell (Figures \ld-e, Ub) with 4.0 to 4.25
moderately rounded, unshouldered whorls. Sexual dimorphism
not significant. Aperture usually separate from body whorl.
Umbilicus open.
Head-foot usually darkly pigmented throughout, except for
tentacles and a broad central patch on sides of head-foot, which
are either lightly dusted or unpigmented.
Ctenidium relatively small (19% of body length), with 16
to 21 filaments, Radula (Figure \2d-f) with relatively few
cusps on lateral teeth and numerous cusps on outer marginals.
Testis large, filling 54% of body length. Penis (Figure 15)
stout. Penial filament extending beyond tip of penial lobe,
width of filament/length, 55%. Glandular ridges, one to three.
Capsule gland (Figure \0d) small (76%) relative to albumen
gland. Seminal receptacle 42% of bursa length.
REMARKS.?This species has a distinctive penis shape and
glandular ridge pattern that is most similar to those of P.
conicus, described next, but differs in having a complex penial
lobe and single, elongate or several circular glandular ridges
on the ventral penial surface.
TYPE-LOCALITY.?Grapevine Spring, Mohave County, Ari-
zona.
DISTRIBUTION.?Grapevine and Whisky Springs in Grand
Wash, north of Lake Mead.
ETYMOLOGY.?Named after Bacchus, the Greek god of wine,
and referring to abundance of wild grapes (Vitis arizonica) in
vicinity of Grand Wash springs.
Pyrgulopsis conicus Hershler, new species
Kingman springsnail
FIGURES lOe, 16a-d, 17, 18
Tryonia new species.?Landye, 1973:42
MATERIAL EXAMINED.?ARIZONA. Mohave County: Burns
Spring, USNM 847208, JJ . Landye and J. King, 20 Apr 1972,
USNM 847215, JJ . Landye, 1 Sep 1972, USNM 847216, JJ .
Landye, 12 May 1980, USNM 847204, R. Hershler and JJ.
Landye, 18 Aug 1985; Dripping Springs, USNM 859039
(holotype), 859040 (3 paratypes), 847237, J J . Landye, 9 May
1980; Cool Spring, USNM 847201,6 Apr 1981.
DIAGNOSIS.?A moderate size species (shell height, 1.8 to
2.7 mm), having an ovate-conic shell. Penis small (15% of
body length), with short filament and single glandular ridge
near tip of penial lobe. Holotype (Figure 16a) height, 2.67
mm; width, 1.69 mm; whorls, 4.25.
DESCRIPTION.?Shells (Figure \6a-d) with 3.5 to 4.5 whorls,
either well rounded with distinct shoulders or only mildly
convex, without shoulders. Sexual dimorphism significant
(females larger than males) in one of two populations studied.
Aperture often loosened from body whorl. Umbilicus open.
Head-foot usually unpigmented, or with light dusting on
snout; subepithelial pigment absent Penial filament pigmenta-
tion light.
Ctenidial filaments, 17 to 21. Central radular teeth (Figure
17) with only four anterior cusps on either side of enlarged,
central cusp. Penis (Figure 18) elongate; filament half as long
as penis, sometimes extending beyond penial lobe. Penial lobe
simple and elongate, with single glandular ridge positioned at
or just proximal to tip of lobe on ventral surface. Less than
half (42%) of bursa posterior to albumen gland (Figure lOe);
seminal receptacle 59% of bursa length.
REMARKS.?The somewhat elongate shell of this species led
to initial assignment of the Burns Spring population to Tryonia
by Landye (1973, 1981).
TYPE-LOCALITY.?Dripping Springs, Mohave County, Ari-
zona.
DISTRIBUTION.?Dripping, Cool, and Burns springs in Black
Mountains near Kingman.
ETYMOLOGY.?From Greek konikos, referring to the conical
shape of shell in this species.
Pyrgulopsis morrisoni Hershler, new species
Page springsnail
FIGURES 10/ \6e-h, 19.20
Fontelicella species.?Williams, 1985:19.
MATERIAL EXAMINED.?ARIZONA. Yavapai County: Tavasci
Springs, USNM 847220, JJ . Landye and D. Belk, 2 Oct 1973;
Page Springs, USNM 859041 (holotype), 859042 (4 para-
types), 847231, JJ . Landye and D. Belk, 2 Oct 1973.
DIAGNOSIS.?A medium size species (shell height, 1.8 to 2.9
mm) with ovate or ovate-conic shell. Penis moderate size, with
narrow filament of medium length, penial lobe relatively large.
Single glandular ridge positioned on ventral surface near tip
of penial lobe. Holotype (Figure \6e) height, 2.00 mm; width,
1.49 mm; whorls, 4.0
DESCRIPTION.?Shell (Figure \6e-h) with 3.75 to 4.50
slightly convex whorls. Sexual dimorphism significant (fe-
males larger than males) in one of two populations studied.
Inner lip thin and usually adnate to body whorl. Aperture less
22
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 15.?Penial variation of P. bacchus, new species, USNM 847203, Grapevine Spring, Mohave County,
Arizona. (Left = dorsal aspect; right = ventral aspect; screened areas = glandular ridges.)
NUMBER 459 23
FIGURE 16.?P. conicus, new species: a, holotype, USNM 859039, Dripping Springs, Mohave County, Arizona;
b, USNM 847203, same locality; c.d, USNM 847204, Burns Spring, Mohave County, Arizona. P. morrisoni,
new species: e, holotype, USNM 859041, Page Springs, Yavapai County, Arizona;/ USNM 847231, same
locality; g.h, USNM 847220, Tavasci Springs, Yavapai County, Arizona. (Bar = 0.75 mm.)
than half of body whorl height. Umbilicus open.
Pigment either absent from head-foot or consisting of light
to moderate dusting throughout (except tentacles). Penial
filament either pigmented along entire length (sometimes
darkly) or unpigmented.
Ctenidial filaments, 15 to 19. Radula (Figure 19) distin-
guished by large numbers of cusps on central and inner
marginal teeth (Table 2). Testis filling 46% to 54% of body
length. Penis (Figure 20) elongate; filament only rarely
extending beyond enlarged penial lobe. All specimens seen
had sole glandular ridge on ventral surface of lobe. Seminal
receptacle 88% to 105% of bursa length.
REMARKS.?The distinctive penis of this species, with a large
lobe, slender filament, and single glandular ridge, is most
similar to that of P. sancarlosensis (described below), but
differs in lacking glandular ridges on the dorsal surface.
TYPE-LOCALITY.?Page Springs, Yavapai County, Arizona.
DISTRIBUTION.?Page and several nearby (within one km)
springs in Verde Valley. The Tavasci Spring population is
probably extinct.
ETYMOLOGY.?Named after the late Dr. J.P.E. Morrison for
his contributions to study of Hydrobiidae.
Pyrgulopsis montezumensis Hershler, new species
Montezuma Well springsnail
FIGURES 10$, 13a,4 2\a-c, 22, 23
Amnicola palomasensis Pilsbry.?Smith, 1953:9.
Fontelicella species.?Williams et al., 1985:19.
MATERIAL EXAMINED.?ARIZONA. Yavapai County: Monte-
zuma Well (swallet), USNM 859043 (holotype), 859044 (3
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 17.?Radula of P. conicus, new species: a, centrals, bar = 12.0 \im; b, laterals, bar = 8.6 Jim; c, laterals
and inner marginals, bar = 16.2 \un; d outer marginals, bar = 6.0 fim.
NUMBER 459 25
FIGURE 18.?Penial variation 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. (Left = dorsal aspect; right = ventral aspect; screened areas = glandular ridges.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 19.?Radula of P. morrisoni, new species: a, centrals, laterals, inner marginals, bar = 13.6 pm; b, centrals,
bar = 12.0 \im; c, inner marginals, bar = 13.6 Jim; d, inner and outer marginals, bar = 23.1 pm.
NUMBER 459 27
mm
FIGURE 20.?Penial variation of P. morrisoni, new species: a,b, USNM 847231, Page Springs, Yavapai County,
Arizona; c4, USNM 847220, Tavasci Springs, Yavapai County, Arizona. (Left = dorsal aspect; right = ventral
aspect; screened areas = glandular ridges.)
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 21.?P. montezumensis, new species: a, USNM 859043, holotype, Montezuma Well, Yavapai County,
Arizona; b,c, USNM 847233, same locality. P. solus, new species: d, USNM 859045, holotype, Brown Spring,
Yavapai County, Arizona; ef, USNM 850290, same locality. P. simplex, new species: g, USNM 859049,
holotype, spring near Strawberry, Gila County, Arizona; h, USNM 847236, locality as above; ij, USNM 847222,
Fossil Springs, Yavapai County, Arizona. (Bar = 0.75 mm.)
paratypes), 847233, J J. Landye and G. Edwards, 10 May 1970;
Montezuma Well (canal at outflow), USNM 847230, JJ .
Landye, 10 May 1970.
DIAGNOSIS.?A moderate size species (shell height, 1.7 to
2.7 mm). Shell ovate to ovate-conic, with slightly loosened
whorls. Distinctive dark pigmentation on snout. Penis large
(29% of body length). Penial filament short, lobe enlarged.
Dorsal penial surface with two elongate glandular ridges (one
on base of filament), one or two additional ridges present on
tip of lobe and on ventral surface. Holotype (Figure 21a)
height, 2.65 mm; width, 1.78 mm; whorls, 4.5.
DESCRIPTION.?Shell (Figures 14a, 21o-c) with 3.5 to 4.5
highly convex whorls, with indented sutures. Loosening of
whorls along axis noticeable in most specimens. Body whorl
73% to 74% of shell height. Sexual dimorphism significant
(females larger than males) in one of two populations studied.
Aperture relatively wide. Inner lip thin, and either just touching
or with much of length fused to body whorl. Umbilicus open.
Operculum (Figure I4d) ovate.
Entire snout, distal half of snout, or just distal lips with very
dark, purple melanin. Tentacles, sides of head/foot, anterior
edge of foot with somewhat lighter pigment. Penial filament
unpigmented or with small pigment patch near base.
Ctenidial filaments, 15 to 19. Lateral angles on central tooth
widely diverging (Figure 22). Central and marginal teeth with
large numbers of cusps (Table 2). Penis (Figure 23) moderately
elongate. Filament only 30% of penial length, rarely extending
beyond penial lobe. Width of filament 40% of length.
Glandular ridges, two to four. Two elongate ridges on dorsal
surface positioned on or just proximal to filament, and near
center of penis. In addition to elongate ridge along (ventral)
edge of lobe, small ridge sometimes present on central portion
of ventral surface. Seminal receptacle 64% of bursa length
(Figure lOg).
REMARKS.?The characteristic slight loosening of shell
coiling in this species is unique among Arizona Pyrgulopsis.
TYPE-LOCALITY.?Montezuma Well, Yavapai County, Ari-
zona.
NUMBER 459 29
FIGURE 22.?Radula of P. monlezumensis, new species, USNM 847233, Montezuma Well, Yavapai County,
Arizona: a, centrals, bar = 12.0 ]im; b, laterals, bar = 8.6 |im; c, laterals and inner marginals, bar = 16.2 fim; d,
outer marginals, bar = 4.3 |im.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
-\ 0.5 mm
FIGURE 23.?Penial variation of P. montezumensis, new species: a,b, USNM 847233, Montezuma Well, Yavapai
County, Arizona; c, USNM 847230, locality as above (but from spring outflow). (Left = dorsal aspect; right =
ventral aspect; screened areas = glandular ridges.)
DISTRIBUTION.?Endemic to Montezuma Well and its upper
100 m of outflow, Montezuma Well National Monument,
Verde Valley.
ETYMOLOGY.?Referring to the type-locality.
Pyrgulopsis solus Hershler, new species
Brown springsnail
FIGURES 21d-/ 24, 25,26a
MATERIAL EXAMINED.?ARIZONA. Yavapai County: Brown
Spring, USNM 859045 (holotype), 859046 (3 paratypes),
850290, J J . Landye, 4 Feb 1986.
DIAGNOSIS.?Shell ovate, small (shell height, 1.4 to
2.0 mm). Penis small (12% of body length), with enlarged,
bifurcate lobe. Elongate glandular ridge along lobe's edge,
smaller ridge on ventral surface. Holotype (Figure 2ld) height,
1.69 mm; width, 1.29 mm; whorls, 3.5.
DESCRIPTION.?Shell (Figure 2\d-f) with 3.5 to 4.0 moder-
ately convex whorls. Body whorl relatively large (83% of shell
height). Sexual dimorphism significant (females larger than
males). Inner lip moderately thickened, fused to or slightly
separated from body whorl. Umbilicus open.
NUMBER 459 31
FIGURE 24.?Radula of P. solus, new species, USNM 850290, Brown Spring, Yavapai County, Arizona: a,
centrals, bar = 8.6 Jim; b, laterals and inner marginals, bar = 7.5 (Jm; c, inner marginals, bar = 6.0 fim; d, outer
marginals, bar = 6.0 \im.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
\ 0.5 mm
FIGURE 25.?Penial variation of P. solus, new species, USNM 850290, Brown
Spring, Yavapai County, Arizona. (Left - dorsal aspect; right - ventral aspect;
screened areas = glandular ridges.)
Pigment on head/foot may be absent, or range from very
light to dark dusting on all surfaces. Penial filament darkly
pigmentcd.
Ctenidial filaments few, 11 to 14. Radula shown in Figure
24. Penis (Figure 25) almost square in shape and very small,
not extending beyond mantle collar. Filament positioned at
only two-thirds of penis length, moderate in length (46% of
penis length), and rarely extending beyond penial lobe. Width
of filament/length, 40%. Bursa (Figure 26a) only 36% of
albumen gland length. Seminal receptacle 83% of bursa length.
REMARKS.?No close relationship is apparent between P.
solus and other Arizona congeners as the minute, squat penis
of this species is unique among southwestern Pyrgulopsis.
TYPE-LOCALITY.?Brown Spring, in southern end of Verde
Valley, Yavapai County, Arizona.
DISTRIBUTION.?Endemic to type-locality.
ETYMOLOGY.?From Latin solus, meaning alone or single,
and referring to endemism of species in a single spring.
Pyrgulopsis simplex Hershler, new species
Fossil springsnail
FIGURES 2\g-j, 26b, 27, 28
Fontelicella species.?Williams et al., 1985:19.
MATERIAL EXAMINED.?ARIZONA. Yavapai County: Fossil
Springs, USNM 847222, J J. Landye, 5 Apr 1972. Gila County:
spring near Strawberry, USNM 859049 (holotype), 859050 (4
paratypes), 847236, J J . Landye, 16 Oct 1971.
DIAGNOSIS.?A moderate size species (shell height, 2.0 to
2.5 mm) with ovate-conic shell. Penis moderate size, with
narrow filament of moderate length and single glandular ridge
on ventral surface of penial lobe. Holotype (Figure 21g) height,
2.41 mm; width, 1.49 mm; whorls, 4.5.
DESCRIPTION.?Shell (Figure 21 g-j) with 3.5 to 4.25
unshouldered and moderately convex whorls. Shell relatively
elongate for genus (shell height/shell width, 150%), with some
specimens conical in shape. Sexual dimorphism significant
(females larger than males) in one of two populations studied.
Aperture usually separate from body whorl. Umbilicus open.
Snout unpigmented, sides of head/foot with light dusting of
melanin. Penial filament darkly pigmented along virtual
entirety of length.
Ctenidium relatively large (27% to 31% of body length),
with 16 to 19 filaments. Radula shown in Figure 27. Penis
(Figure 28) very elongate. Filament 57% of penis length,
projecting well beyond tip of penial lobe. Width of filament
only 28% of length. Variably sized, single, glandular ridge
positioned near distal edge of simple penial lobe. Ovary only
12% of body length. Over half (56%) of bursa posterior to
albumen gland. Seminal receptacle 65% of bursa length.
REMARKS.?P. simplex may be closely allied with other
Arizona species having a relatively simple penis, including P.
conicus, P. morrisoni, and P. sancarlosensis. The penial
filament of P. simplex is shorter than that of P. sancarlosensis,
but longer than those of the other two species.
TYPE-LOCALITY.?Spring near Strawberry, Gila County,
Arizona.
DISTRIBUTION.?Restricted to a number of springs forming
the perennial portion of Fossil Creek, which flows into Verde
River south of Verde Valley.
ETYMOLOGY.?From Latin simplex, referring to simple
condition of penis, with few glandular ridges.
Pyrgulopsis confluentis Hershler, new species
Three Forks springsnail
FIGURES 3d-g, 26c, 29, 30
New genus, new species.?Landye, 1973:49.
New genus, new species.?Landye, 1981:27.
MATERIAL EXAMINED.?ARIZONA. Apache County: spring
on N side of Blanket Creek at Three Forks, USNM 859053
(holotype), 859054 (4 paratypes), 847234, JJ . Landye, 15 Sep
1977; spring at Three Forks, USNM 847229, J J . Landye, 7
Oct 1973.
DIAGNOSIS.?Shell low ovate to ovate-conic, variable in size
(shell height, 1.5 to 4.8 mm). Penis moderate size, with small
filament; elongate, tapering lobe; and enlarged ventral
accessory crest. Glandular ridges numerous on both penial
surfaces. Holotype (Figure 3d) height, 2.80 mm; width, 1.96
mm; whorls, 4.5.
DESCRIPTION.?Shell with 3.5 to 4.75 unshouldered and
FIGURE 26.?Pallial oviducts of Pyrgulopsis species: 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 of Blanket 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; / P. thompsoni, new species, USNM 847238, Peterson Ranch Springs, Santa Cruz County,
Arizona.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 27.?Radula of P. simplex, new species, USNM 847236, spring near Strawberry, Gila County, Arizona:
a, centrals, bar = 12.0 \un, b, laterals, bar = 13.6 ^m; c, inner marginals, bar = 10.0 \im; d, outer marginals,
bar = 7.5 ujn.
NUMBER 459 35
^ 0.5 mm
FIGURE 28.?Penial variation of P. simplex, new species: a, b, USNM 847236, spring near Strawberry, Gila
County, Arizona; c, USNM 847222, Fossil Springs, Yavapai County, Arizona. (Left = dorsal aspect; right =
ventral aspect; screened areas = glandular ridges.)
well-rounded whorls. Note shell size and shape difference
among the two known populations (Figure 3d-g). Sexual
dimorphism significant (females larger than males) in one
population. Dark, thickened periostracum covering entire shell
surface. Inner lip thin. Umbilicus chink-like to open.
Dark pigment on snout and tentacles characteristic, pigment
somewhat lighter on sides of head/foot. Proximal half of penial
filament darkly pigmented, with pigment sometimes extending
into penis as well.
Ctenidial filaments, 17 to 23. Ctenidium only 19% to 21%
of body length. Radula shown in Figure 29. Inner marginals
with as many or more cusps as outers (Table 2). Testis 46%
of body length. Penis elongate (Figure 30); filament only 27%
of penis 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 swollen. Glandular ridges, four to six, rarely
enlarged or elongate. One ridge (sometimes absent) on
proximal (dorsal) half of filament; two on dorsal portion of
lobe (or just proximal to lobe); one just proximal to tip of lobe
(ventral); and one or two on accessory crest. Ovary only 11%
of body length. Oviduct coil complex, coil diameter only 30%
of albumen gland length (Figure 26c). Half of bursa posterior
to albumen gland. Seminal receptacle 70% of bursa length.
REMARKS.?This species links with the P. glandulosus-
group on the basis of penial morphology, but differs in lacking
accessory crests.
TYPE-LOCALITY.?Spring on N side of Blanket Creek at
Three Forks, Apache County, Arizona.
DISTRIBUTION.?Several springs at Three Forks feeding
Black River.
ETYMOLOGY.?From Latin confluens, meaning a place where
streams meet, and referring to occurrence of species at Three
Forks.
Pyrgulopsis sancarlosensis Hershler, new species
Bylas springsnail
FIGURES \3e, 26e, 3\a-g, 32, 33
Fonlilkella [sic] species.?Meffe and Marsh, 1983:369.
Fontelicella species.?Williams et al., 1985:19.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 29.?Radula of P. confluenlis, new species, USNM 847234, spring on N side of Blanket Creek at Three
Forks, Apache County, Arizona: a, centrals, bar = 12.0 Jim; b, laterals and inner marginals, bar = 12.0 (Am; c,
laterals and inner marginals, bar = 13.6 |im; d, outer marginals, bar = 7.5 |im.
NUMBER 459 37
FIGURE 30.?Penial variation 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. (Left = dorsal aspect; right = ventral aspect; screened areas = glandular ridges.)
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
h
FIGURE 31.?P. sancarlosensis, new species: a, USNM 859051, holoiype, springs W of Tom Niece Springs,
Graham County, Arizona; b,c, same locality, USNM 847226; d,e, USNM 847224, large spring N of Bylas,
Graham County, Arizona; f,g, USNM 847217, small spring near Bylas, Graham County, Arizona. P. cochisi,
new species: h, holotype, USNM 859055, springs at San Bernardino Ranch, Cochise County, Arizona; i-k,
same locality, USNM 847218. (Bar = 0.75 mm.)
NUMBER 459 39
FIGURE 32.?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 \isn\ c, laterals and inner marginals, bar = 8.6 fim; d,
laterals, bar = 15.0 |im; e, inner marginals, bar = 6.0 \im\f, outer marginals, bar = 8.6 \im.
MATERIAL EXAMINED.?ARIZONA. Graham County: small
spring near Bylas, USNM 847217, J.J. Landye, 20 May 1971;
spring near Gila River bridge NW of Bylas, USNM 847213,
J.J. Landye, 30 Jan 1971; large spring N of Bylas, USNM
847224, J.J. Landye and D.W. Taylor, 20 Apr 1971; springs
W of Tom Niece Springs, USNM 859051 (holotype), 859052
(4 paratypes), 847226, JJ . Landye, 29 Jun 1973; seeps W of
Tom Niece Springs, USNM 847225, J J . Landye, 27 Jun 1973.
DIAGNOSIS.?Shell small (height, 1.1 to 2.4 mm), varying
in shape from globose to elongate-conic. Penis large (25% to
33% of body length), with elongate filament and up to three
glandular ridges. Seminal receptacle absent. Holotype (Figure
31a) height, 1.90 mm; width, 1.29 mm; whorls, 4.0.
DESCRIPTION.?Shell height, 1.1 to 2.4 mm; whorls, 3.25 to
4.25, moderately rounded. Diversity in shell size and shape
shown in Figure 31a-g. Both shell forms sometimes present
in single population, with conical specimens usually enlarged
(often female). Sexual dimorphism not significant. Small
globose specimens with relatively large apertures (50% to 55%
of shell height); larger, conical shells with smaller apertures
(48% of shell height). Aperture usually separated, sometimes
greatly so, from body whorl. Umbilicus open. Operculum
shown in Figure 13e.
Snout darkly (black) pigmented, tentacles and sides of
head/foot with lighter pigment, central section of latter
sometimes unpigmented. Penial filament either darkly pig-
mented along entire length or unpigmented.
Ctenidial filaments, 12 to 20. Radula shown in Figure 32.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 33.?Penial variation 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.
(Left - dorsal aspect; right = ventral aspect; screened areas = glandular ridges.)
NUMBER 459 41
Penial filament relatively narrow (width/length, 26% to 29%)
and elongate (67% to 84% of penis length), but usually
projecting perpendicular to penis length and therefore not
extending beyond tip of penial lobe. Enlarged lobe usually
untapered. Glandular ridges, one to three. Dorsal surface of
penis with or without single ridge; ventral surface with single
enlarged ridge and sometimes an additional smaller one.
Capsule gland (Figure 26e) consistently longer than albumen
gland. Oviduct coil diameter large (81% to 82%) relative to
albumen gland length. Bursa 60% to 99% of albumen gland
length. Note that duct exits lateral to bursa tip.
REMARKS.?Absence of seminal receptacle is not known for
any congener, although size of structure does vary greatly and
is greatly reduced in forms such as P. cochisi (described next).
Loss of seminal receptacle may represent the end point of a
reduction trend, with sperm storage function presumably
assumed by the thickened coiled section of oviduct.
TYPE-LOCALITY.?Springs W of Tom Niece Springs, Graham
County, Arizona.
DISTRIBUTION.?Springs along north side of Gila River
between By las and Pima.
ETYMOLOGY.?Named after San Carlos Apache Indian
Reservation near Bylas.
Pyrgulopsis cochisi Hershler, new species
San Bernardino springsnail
FIGURES 254 Wh-k, 33, 34
Fontelicella species.?Williams etal., 1985:50.
MATERIAL EXAMINED.?ARIZONA. Cochise County: Springs
at San Bernardino Ranch, USNM 859055 (holotype), 859056
(2 paratypes), 847218, J.J. Landye, 20 Mar 1971.
DIAGNOSIS.?A small size species (1.3 to 1.7 mm) having
ovate-conic shell. Penis moderate size, without lobe; filament
elongate. Dorsal and ventral penial surfaces each with single
glandular ridge. Holotype (Figure 31 h) height, 1.55 mm; width,
1.04 mm; whorls, 3.5.
DESCRIPTION.?Shell (Figure 32h-k) with 3.25 to 4.0 slightly
convex whorls, with sutures only very slightly indented. Sexual
dimorphism significant (females larger than males). Inner lip
thin. Umbilicus open.
Snout with light to dark dusting of pigment. Sides of
head/foot somewhat less pigmented than snout, although
operculigerous lobe often quite dark. Penial filament darkly
pigmented along entire length.
Ctenidial filaments, 12 to 15. Anterior portion of central
tooth with few cusps (4-14) (Figure 34). Inner and outer
marginals with roughly same number of cusps (Table 2).
Prostate 11% of body length. Penis fairly elongate (Figure
35). Penial filament 61% of penis length, extending distal to
anterior tip of penis. Filament three times as long as wide.
Glandular ridges, two. 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% of bursa length.
REMARKS.?Pyrgulopsis cochisi is most similar to P.
thompsoni, which also have small penes with reduced penial
lobe (absent in the former) and single glandular ridge, but is
smaller and has a glandular ridge on both penial surfaces.
TYPE-LOCALITY.?Spring at San Bernardino Ranch, Cochise
County, Arizona.
DISTRIBUTION.?Restricted to type-locality and nearby small
seeps at San Bernardino Ranch.
ETYMOLOGY.?Named after Apache chief Cochise.
Pyrgulopsis thompsoni Hershler, new species
Huachuca springsnail
FIGURES 26/ 36,37, 38
Paludestrina stearnsiana.?Pilsbry and Ferriss, 1909:516.
"Undescribed...hydrobiid."?Minckley, 1969:6.
Fontelicella species.?Bequaert and Miller, 1973:214.
Fontelicella species.?Williams et al., 1985:19.
MATERIAL EXAMINED.?ARIZONA. Santa Cruz County: Cot-
tonwood Springs, USNM 847221, J.J. Landye, 19 Aug 1973;
Monkey Spring, USNM 847223, JJ . Landye and D. Belk, 2
Dec 1969; Canelo Hills Cienega, USNM 847228, J J. Landye,
14 Nov 1978; Sheehy Spring, USNM 847235, J.J. Landye, 18
Apr 1976, USNM 847227, 14 Nov 1978; Peterson Ranch
Springs, USNM 859057 (holotype), 859058 (3 paratypes),
847238, JJ . Landye, 19 Aug 1973. SONORA (Mexico): Ojo
Caliente, USNM 847251, JJ . Landye and D.W. Taylor, 7 Apr
1971.
DIAGNOSIS.?A moderate to large size species (1.7 to 3.2
mm) having ovate or ovate-conic shell. Penis moderate size,
with elongate filament and single, small glandular ridge on
ventral surface of penial lobe. Holotype (Figure 36a) height,
2.57 mm; width, 1.63 mm; whorls, 4.25.
DESCRIPTION.?Shell (Figure 36) with 3.25 to 5.0 moderately
convex, slightly shouldered whorls. Sexual dimorphism
significant in two of four populations studied, with males larger
in one of these and females larger in other. Inner lip thin.
Aperture fused to or separate from body whorl. Umbilicus
chink-like or open.
Snout and anterior portion of foot (dorsal surface) with light
to fairly dark pigment. Remainder of head/foot usually
unpigmented. Penial filament darkly pigmented throughout
Ctenidial filaments, 11 to 20. Radula shown in Figure 37.
Testis 37% to 54% of body length. Prostate 7% to 8% of body
length. Penis squat to elongate. Penis filament centered at 80%
to 93% of penis length. Filament 35% to 103% of penis length,
and only slightly thickened (width/length, 27% to 46%). Penis
lobe simple. Glandular ridge usually located near tip of lobe.
Oviduct coil relatively complex in that initial turn of loop bends
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 34?Radula of P. cochisi, new species, USNM 847218, springs at San Bernardino Ranch, Cochise
County, Arizona: a, centrals, bar = 9.1 jun; b, laterals, bar = 5.0 nm; c, laterals and inner marginals,
bar = 6.7 Jim; d, outer marginals, bar = 6.0 Jim.
NUMBER 459 43
-\ 0.5 mm
FIGURE 35.?Penial variation of P. cochisi, new species, USNM 847218,
springs at San Bernardino Ranch, Cochise County, Arizona. (Left = dorsal
aspect; right = ventral aspect; screened areas = glandular ridges.)
back on itself (Figure 26/). Most of bursa length (55% to 85%)
posterior to albumen gland. Seminal receptacle 53% to 72%
of bursa length.
TYPE-LOCALITY.?Peterson Ranch Springs, Santa Cruz
County, Arizona.
DISTRIBUTION.?Six springs in Southern Santa Cruz County,
Arizona, United States, and northern Sonora, Mexico.
ETYMOLOGY.?Named after Dr. F.G. Thompson, in recogni-
tion of his contributions to study of North American
Hydrobiidae.
Genus Tryonia Stimpson, 1865
DIAGNOSIS.?Shell elongate-conic to turreted, 1.7 to 7.0
mm high. Aperture simple, umbilicus narrow or absent
Teleoconch smooth or with spiral lines or collabral striations
or varices. Penis with enlarged glandular lobe at base and one
to four smaller glandular lobes on inner curvature. Females
ovoviviparous, with young brooded in capsule gland. Thin-
walled capsule gland reflected posteriorly, albumen gland
greatly reduced in size. Bursa copulatrix and seminal receptacle
small (relative to pallial oviduct), spermathecal duct short.
REMARKS.?Hershler and Thompson (1987) provided a
detailed description for the genus. Hyalopyrgus Thompson,
1968 is regarded by them to be a synonym of Tryonia. The
genus has a disjunct distribution, occurring in much of Florida
as well as a large part of the arid Southwest. These snails are
related to a group of genera, including Aphaostracan
Thompson, 1968, Littoridinops Pilsbry, 1952, Mexipyrgus
Taylor, 1966, and Pyrgophorus Ancey, 1888 that have
elongate-conic shells and glandular (mammiform) penial lobes.
Tryonia differs from the above by its turreted shell and unique
position of penial lobes. The genus contains ten described
species as well as numerous undescribed species from the
Southwest.
Tryonia gilae Hershler, new species
Gila tryonia
FIGURES 1 4 C / 39a-e, 40, 41a-c, 42,43d-i, 44, 45, 46a, 4,1b
Tryonia species.?Meffe and Marsh, 1983:369.
Tryonia species.?Williams et al., 1985:19.
MATERIAL EXAMINED.?ARIZONA. Graham County: small
spring near Bylas, USNM 859059 (holotype), 859060 (4
paratypes), 847257, J J . Landye, 30 Jan 1971; spring near Gila
River bridge NW of Bylas, USNM 847255, J.J. Landye, 30 Jan
1971; large spring N of Bylas, USNM 847258, J J . Landye and
D.W. Taylor, 20 May 1971; springs W of Tom Niece Springs,
USNM 847254, JJ . Landye, 27 Jun 1973; seeps W of Tom
Niece Springs, USNM 847252, JJ . Landye, 27 Jan 1973; Cold
Springs, USNM 847253, JJ . Landye, 27 Jan 1973.
DIAGNOSIS.?A moderately large size species (1.9 to
3.3 mm) with elongate-conic to turreted shell. Cephalic
tentacles with well-defined linear tracts of hypertrophied cilia.
Penis with two glandular lobes on inner curvature near distal
tip and enlarged lobe on outer curvature at base. Outer
curvature distal to lobes with hypertrophied ciliary tuft. Bursa
relatively large. Holotype (Figure 39a) height, 3.25 mm; width,
1.45 mm; whorls, 6.0.
DESCRIPTION.?Shell (Figure ?>9a-e) clear, transparent, and
without periostracum. Shell often twice as tall as wide; whorls,
4.5 to 6.0, slightly to moderately convex with indented sutures.
Sexual dimorphism pronounced, with females larger than
males. Protoconch (Figure 14c) smooth and flat (sometimes
slightly depressed), with 1.0 to 1.25 whorls. Embryonic shell
with as many as 1.5 whorls. Body whorl 57% to 74% of shell
height. Aperture simple, occupying less than half of shell
height, longer than wide, slightly angled above and rounded
below, and either fused to or separate from body whorl. Inner
lip fairly straight, slightly thickened and reflected; outer lip
rounded and thin. Apertural plane slightly tilted from coiling
axis (somewhat advanced adapically) with long axis inclined
25? to 30? to coiling axis. Umbilicus open. Teleoconch with
strong growth lines. Operculum (Figure 14/) amber, paucispi-
ral, over one and a half times longer than wide, and with three
whorls.
Snout (Figure 40) longer than wide, terminating with fleshy
lips. Cephalic tentacles (Figures 40, 4la,b) narrow, slightly
expanded at tips, and moderately elongate (slightly extending
beyond snout tip). Ciliary tracts on tentacles shown in Figure
41a-c. Snout and "neck" with scattered ciliary tufts. Black
eyes located in slightly bulging outer edges of tentacle bases.
Foot elongate, with anterior edge almost straight and posterior
portion expanded and rounded. Anterior pedal glands, 12 to 18.
Head/foot lightly dusted with epithelial melanin throughout,
except for tentacles. Narrow strip extending down sides 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
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 36.?P. thompsoni, new species: a, holotype, USNM 859057, Peterson Ranch Springs, Santa Cruz
County, Arizona; b, USNM 847238, same locality; cA USNM 847221, Cottonwood Springs, Santa Cruz
County, Arizona; ej, USNM 847228. Canelo Hills Cienega, Santas Cruz County, Arizona; g.h, USNM 847223,
Monkey Springs, Santa Cruz County, Arizona; i-k, USNM 847227, Sheehy Spring, Santa Cruz County, Arizona;
/, USNM 847251, Ojo Caliente, Sonora, Mexico. (Bar = 0.75 mm.)
FIGURE 37.?Radula of P. ihompsoni, 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; ej. laterals and inner marginals, USNM 847238. bars = 6.7,
7.5 um; g, inner marginals, USNM 847238, bar = 10.0 um; M, outer marginals, USNM 847238, bars = 6.0 um.
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
0.5 mm
FIGURE 38.?Penial variation 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;/ USNM 847223, Monkey Spring, Santa Cruz
County, Arizona; g, USNM 847227, Sheehy Spring, Santa Cruz County, Arizona; h, USNM 847251, Ojo
Caliente, Sonora, Mexico. (Left = dorsal aspect; right = ventral aspect; screened areas = glandular ridges.)
NUMBER 459 47
FIGURE 39.?Tryonia gilae, new species: a, holotype, USNM 859059, small spring near Bylas, Graham County,
Arizona; b,c, same locality, USNM 847257; d,e, USNM 847253, Cold Springs, Graham County, Arizona. T.
quitobaquitae, new species:/ holotype, USNM 859061, Quitobaquito Springs, Pima County, Arizona; g.h,
same locality, USNM 847256. (Bar = 0.75 mm.)
pallial cavity roof above ctenidium darkly pigmented in males.
Penis with dark pigment patch near distal tip (see Hershler,
1985, fig. 44) and smaller pigment streaks at bases of small
penial lobes and elsewhere on dorsal surface. Dark subepithe-
lial pigment concentrated on right side of mantle collar. Large,
dull grey glandular clusters in "neck" proximal to tentacles.
Bases of tentacles with white glandular clusters.
Ctenidium (Figure 42, Ct) elongate, filling most of pallial
cavity. Ctenidial filaments, 20 to 25. Filaments broad.
Osphradium (Figure 42, Os) small relative to ctenidium length.
Kidney (Figure 44a, Ki) with small opening fringed by obvious
lips (Figure 44a, Oki).
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Cgp Sts
 a Acg
FIGURE 40.?Dorsal aspect (without shell) of relaxed T. gilae, new species, USNM 847257, small spring near
By las, Graham County, Arizona: a, female; b, male. (Abbreviations: Acg = anterior end of capsule gland;
Cgp = posterior end of capsule gland; Ct = ctenidium; Em = brooded embryo; In = intestine; Op = operculum;
Pc = pericardium; Pn = penis; Sts = style sac.)
Buccal mass small (about 13%) relative to snout. Radula
(Figure 43d-i) typically taenioglossate, with broadly trapezoi-
dal central teeth having two pairs of basal cusps. Lateral angles
of centrals widely divergent, narrow basal process character-
istic. Central teeth with enlarged central cusp; central cusp of
laterals sometimes not enlarged (Figure 43g). Inner and outer
marginals similarly shaped; outers with only a few more cusps
than inners. Stomach (Figure 44b, Ast, Pst) and style sac (Sts)
about equal in length; 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 half of stomach and
extending (ventral to testis) almost to tip of visceral coil.
Testis large, overlapping most of posterior stomach cham-
ber. Prostate gland (Figure 44a, Pr) small, flat, near-circular
in outline, positioned largely or totally posterior to pallial
cavity. Vas deferens enters posterior end of prostate gland and
exits near anterior tip.
Penis (Figure 40, Pn) flattened, elongate, large relative to
snout, extending forward from attachment area without coiling.
Two small penial lobes (Figure 46a, PI) on inner curvature
centered at about two-thirds of penis length. Lobe on outer
curvature massive, and usually folded back against penis near
base. All penial lobes bearing mammiform glands distally. Tip
of penis with pronounced swelling on inner curvature (Figure
46a, Ics). Vas deferens undulating in penis, terminating in
eversible papilla lateral to swelling (Figure 46a, Vd). Penis
surface, especially at base, well endowed with dense glandular
clusters of several types. Penial ciliation shown in Figure 45.
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 (Figure 45a,b). Three or four distinctive
strips of cilia cross penis surface proximal to enlarged lobe
(Figure 45c,<f). Remainder of penis surface with scattered,
small ciliary tufts.
Ovary very small relative to body length, non-lobular.
Capsule gland (Figure 40a, Cg) enlarged (covering entirety of
style sac), containing three or four shelled embryos. Brooded
young darkly pigmented. Capsule gland with narrow muscular
sphincter (Figure 42, Cg) at anterior end as well as posterior
bend (Figure 47b, Cg) which extends (partly ventral to distal
section of capsule gland) almost to anterior end of pallial
cavity. Albumen gland (Figure 47c, Ag) comprising very
small, narrow section of pallial oviduct, terminating ventral to
NUMBER 459 49
FIGURE 41.?SEM micrographs of cephalic tentacles oiTryonia gilae, new species, USNM 847257, small spring
near Bylas, Graham County, Arizona: a, left tentacle, bar - 100 pm; b, right tentacle, bar = 120 \xm; c, close-up
of ciliation on tentacle, bar = 20.0 \im. T. quitobaquitae, new species: d, left tentacle, bar = 75 pm; e, right
tentacle, bar = 75 Jim;/ distal half of penis, showing dense ciliation, bar = 50 (im.
bursa copulatrix (Bu) at entry of renal oviduct. Oviduct (Ov)
with simple coil (ventral to bursa) proximal to opening into
albumen gland (Oov). Bursa club-shaped, small relative to
pallial oviduct, and largely obscured anteriorly by albumen
gland. Bursa gradually narrows anteriorly to give rise to
spcrmathecal duct (Sd) just posterior to end of pallial cavity.
Seminal receptacle (Sr) minute, with short duct connecting to
oviduct. After this juncture duct (Dsr) coils several times
ventral to bursa before looping back to open into dorsal surface
of bursa at or just proximal to origin of spcrmathecal duct
(Osr). Spermathecal duct pressed to ventral surface
(columellar side) of capsule gland. Duct narrow, non-muscular,
and short, opening just anterior to posterior end of pallial
cavity (Figure 42).
REMARKS.?This species is not very similar to either T.
quitobaquitae from Arizona (described next) or undescribed
taxa from New Mexico (Taylor, 1983). Its affinities may lie
instead with Nevada species having relatively many penial
lobes.
TYPE-LOCALITY.?Small spring near Bylas, Graham County,
Arizona.
DISTRIBUTION.?Springs along north side of Gila River
between Bylas and Pi ma.
ETYMOLOGY.?Referring to distribution of this species along
Gila River.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Cg
Emc
FIGURE 42.?Pallial cavity (viewed from right) of T. gilae, new species, USNM 847257, small spring near
Bylas, Graham County, Arizona: Most of capsule gland removed (position indicated by dashed lines) to reveal
ctenidium and spermathecal duct. (Abbreviations: Cg = capsule gland; Q = ctenidium; Emc = posterior end of
pallial cavity; In = intestine; Me = mantle collar, Os = osphradium; Sd = spermathecal duct.)
Tryonia quitobaquitae Hershler, new species
Quitobaquito tryonia
FIGURES 39f-K *\d-f. 43a-c, 46a, 47a
Tryonia imitator (Pilsbry).?Bequaert and Miller, 1973:213.
MATERIAL EXAMINED.?ARIZONA. Pima County: Quitoba-
quito Springs, USNM 859061 (holotype), 859062 (4 para-
types), 847256, J.J. Landye, 13 Mar 1971.
DIAGNOSIS.?A minute size species (1.4 to 1.8 mm), with
ovate-conic to turreted shell. Cephalic tentacles without
well-defined ciliary tracts. Penis with two lobes on inner
curvature, a small one just distal to midpoint and a somewhat
larger one near the base. Much of distal portion of penis with
dense cover of cilia. Bursa very small relative to pallial oviduct
Holotype (Figure 39/) height, 1.96 mm; width, 0.96 mm;
whorls, 4.5.
DESCRIPTION.?Shell (Figure 39/-A) with 3.5 to 4.5 highly
convex whorls with deep sutures. Apertural lip thin. Inner lip
only slightly reflected and usually fused to body whorl.
Umbilicus chink-like.
Dark pigmentation on visceral coil near uniform. Snout and
operculigerous lobe usually darkly pigmented, elsewhere
head/foot having light dusting of melanin, with broad central
section on sides of head/foot sometimes having distinctively
light coating. Penial tip with pigment patch.
Scattered ciliary tufts on cephalic tentacles shown in Figure
4\a\e. Central teeth of radula (Figure 43a) with single pair of
basal cusps. Lateral teeth (Figure 436) with enlarged central
cusp. Marginals shown in Figure 43b,c.
Elongate penis shown in Figure 46b. Note that vas deferens
undulates only slightly in penis. Extensive penial ciliation
shown in Figure 41/. Ciliation extending on both outer and
inner curvatures to level of base of distal lobe. Bursa copulatrix
complex shown in Figure 47a,b. Bursa (Bu) minute, not greatly
overlapped by albumen gland.
REMARKS.?The reduced lobation of T. quitobaquitae is
shared by two large undescribed congeners from New Mexico
(Taylor, 1983).
TYPE-LOCALITY.?Quitobaquito Springs, Organ Pipe Na-
tional Monument, Pima County, Arizona.
DISTRIBUTION.?Type-locality, Burro Spring and an un-
named spring, all within the national monument.
ETYMOLOGY.?Referring to the type-locality.
Morphometrics
Results of discriminant analyses based on shell and
anatomical data are in Tables 3-5, with scores on the first two
discriminant functions plotted in Figures 48 and 49. Correct
classification to species for grouped cases, representing
topotypes for each species, was 89% (overall) for the analysis
based on male shell data (ranging from 80% to 100% among
groups), 93% for the analysis based on female shell data (63%
to 100%), and 100% for analyses using anatomical data (Table
6). Runs using subsets of taxa, such as the Verde Valley
FIGURE 43.?Radula of Tryonia quitobaquitae, new species, USNM 847256, Quitobaquilo Springs, Pima County,
Arizona: a, centrals, bar = 8.6 |im; b, laterals and inner marginals, bar - 6.7 \im; c, outer marginals,
bar = 5.0 \im. T. gilae, new species: d, centrals, USNM 847257, small spring near Bylas, Graham County,
Arizona, bar = 10.0 (im; e, centrals, USNM 847253, Cold Springs, Graham County, Arizona, bar = 7.5 |i.m; /
laterals, USNM 847257, bar = 6.0 yon; g, laterals, USNM 847253, bar = 5.0 \un\ h, inner marginals, USNM
847253, bar = 12.0 ujn; i, outer marginals, USNM 847257, bar = 7.5 urn.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
,Pr
a
Ast
Sts
Sts
FIGURE 44.?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; Emc = posterior end of pallial cavity; In = intestine; Ki = kidney;
Oes = oesophagus; Oki = kidney opening; Pr = prostate gland; Pst = posterior stomach chamber, Sts = style
sac; Ve = ventricle.)
species, did not yield superior classification.
The somewhat lower level of successful classification based
on shell data is reflected in plots of scores on the first two
discriminant functions (Figure 48), which collectively explain
68.8% (male) and 78.7% (female) of total variance. Note that
groups are not well separated, especially in the analysis of
female shell data (Figure 48ft) in which each is overlapped by
at least one other. Significant separation of closest groups (7,
P. simplex; 3, P. conicus) was not achieved even after all steps
were completed (p = 0.33): note that all points representing the
former are enclosed within the latter. For the analysis of male
data (Figure 48ft), significant separation of closest groups was
achieved (p = 0.04), yet all except two exhibit overlap.
Interpretation of the two discriminant functions (based on
loadings and correlations in Table 3) suggests that function 1
represents size in both (decreasing along axis in male plot,
increasing in female plot) while function 2 correlates with
relative shell height (increasing along axis in male plot,
decreasing in female plot). Note that most separation of groups
occurs along the size-correlated axis. The two non-overlapped
groups in the male plot (\,P. deserta, separated on axis II; 12,
P. cochisi, separated on axis I) are unusually globose-shelled
and small, respectively. Classification of ungrouped cases was
highly variable, ranging from 0% for several populations oiP.
thompsoni to 70% for an additional population of that
species.
A superior classification resulting from analyses based on
anatomical data is reflected in relatively separated groups in
plots of scores on the first two discriminant functions
(collectively explaining 64% (male) and 67.6% (female) of
total variance), with six (male) and three (female) groups
non-overlapped (Figure 49a,ft). Significant (p>.005) separation
of closest groups was achieved after the final step (in both).
In the plot based on analysis of male anatomical data,
separation of groups is fairly good along both axes (especially
1), and largely interpretable in terms of penial morphology and
body size. Function 1 has particularly high loadings of GLR
and WFI (Table 4). Species with low scores on this axis, such
as P. thompsoni (11), have penes with thickened filaments and
few (usually one) glandular ridges (see Figure 38), while those
with higher scores (9, P. confluentis; 8, P. glandulosus; 5, P.
montezumensis) have penes with narrow filaments and more
than two ridges (see Figures 8, 9, 23, 30). The second axis has
high loadings of other penial variables (L2, L3/L2, LFI) as
well as a size component (note high loading of BL). Species
with low scores on this axis have relatively long penial
filaments and/or lobes (Figures 14, 28, 31, 35) (10, P.
sancarlosensis; 7, P. simplex; 1, P. deserta; 12, P. cochisi),
while species with high scores, such as P. bacchus (2), have
shortened filaments and/or lobes (Figure 15). Note that
ungrouped specimens are fairly tightly clustered and positioned
close to topotypes of their species (especially P. sancarlo-
sensis), but rarely overlap with them (Figure 49a). For these
ungrouped specimens, classification ranged from 0% (two
NUMBER 459 53
FIGURE 45. 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 urn; b, close-up of distal end of above, bar = 75 pm; c,d,
dorsal aspect of penis base, bars = 86, 100 \im.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
a
Ics
0.25 mm
FIGURE 46.?Penis of T. gilae, new species: a, (above) USNM 847257, small spring near Bylas, Graham County,
Arizona. T. quitobaquilae, new species: b, Quitobaquito Springs, Pima County, Arizona. (Abbreviations: Ics =
swelling on inner curvature; PI = penial lobe; Vd = vas deferens.
populations of P. thompsoni) to 100% (a third population of
the same).
In the plot based on analysis of female data (Figure 49b),
group separation is largely effected by scores on the first
discriminant function, which is clearly size-correlated, given
high loadings of CTL, OSL, OCT, and BDW on it (Table 5).
The second discriminant function has high loadings of
OSL/CTL, BUW, COP, and SRL/BUL. The particularly low
scores of P. sancarlosensis (10) on this axis probably result
from absence of a seminal receptacle in this species.
Ungroupcd specimens (especially P. sancarlosensis and
P. morrisoni) are generally more separated from
appropriate topotypes than in the male plot. Also note the large
variance on the first discriminant function for specimens
representing P thompsoni. For ungrouped cases, classification
ranged from 0 to 40%.
Discussion
Results of the discriminant analyses support division of
Arizona Pyrgulopsis 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 them.
Discriminant analyses using anatomical data provide classifica-
tions slightly superior to those generated by shell data, and
yield greater species separation on plots (especially based on
male anatomical data) of the first two discriminant function
scores, suggesting the former data set discriminates these
species somewhat better than the latter. This may be partly due
Cg a
Osr
H 0.5 mm
Sts
FIGURE 47.?Bursa copulalrix complex (viewed from right) of T. quitobaquitae, new species: a,b USNM 847256,
Quitobaquito Springs, Pima County, Arizona, and T. gilae, new species: cjd, USNM 847257, small spring near
Bylas, Graham County, Arizona; much of pallial oviduct cut away (position indicated by dashed lines) to reveal
underlying structures; visceral ganglion connective tightly pressed against oviduct. (Abbreviations: Ag = albumen
gland; Bu = bursa copulatrix; Cg = capsule gland; Cvg = visceral ganglion connective; Dsr = duct of seminal
receptacle; Emc = posterior end of pallial cavity; 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; Ov = oviduct; Sd = spermathecal duct; Sr = seminal receptacle; Sts = style sac.)
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
43.4% 56.6%
FIGURES 48,49?Plots of scores on first two discriminant functions derived from Pyrgulopsis shell data (Figure
48, above) and anatomical data (Figure 49, below); percent variation explained by each function is indicated
on respective axes; ellipses encircle positions of topotypical specimens for each species; positions of ungrouped
specimens (from additional populations) for three species are indicated by symbols in Figure 49 (a = male; b -
female; 1 = P. deserta; 2 - P. bacchus; 3 = P. conicus; 4 = P. morrisoni; 5 = P. montezumensis; 6 = P. solus;
7 = P. simplex; 8 = P. glandulosus; 9 = P. confluentis; 10 = P. sancarlosensis; 11 = P. thompsoni; \2 = P.
cochisi.)
21
.6
%
II
11i
A A i
A
7
a
P
1 AA / * / 4
^ Or
10
I
; (
A 4
? 10
42.4%
i
20
.1
II
12
4
(
ft
8 A
5
1
A
A
0
A 4
? 10
46.9%
NUMBER 459 57
TABLE 3.?Results of stepwise discriminant function analysis on thirteen shell variables in terms of standardized
weights and correlation of variable with discriminant function.
Variable
SH
SW
AH
AW
LBW
WBW
CD
AB
NW
SH/SW
AH/SH
LBW/SH
CD/AB
Function 1
0.56
2.38
3.91
-0.21
-6.93
-0.62
-0.69
0.55
-0.12
1.83
-2.52
2.61
0.64
-0.61
-0.72
-0.69
-0.73
-0.69
-0.79
-0.30
-0.55
-0.36
0.06
0.01
0.09
0.08
Males
Function 2
2.44
1.10
5.54
0.46
-5.21
0.01
-0.86
1.11
0.27
1.35
-3.28
1.59
1.04
0.53
0.27
0.36
0.37
0.48
0.40
0.38
0.50
0.48
0.61
-0.54
-0.46
0.10
Females
Function 1
-2.21
1.12
-0.91
-0.08
1.38
0.89
-1.17
0.64
0.13
0.56
0.58
-1.08
0.83
0.51
0.82
-3.57
0.68
0.72
0.82
0.31
0.55
0.33
0.08
-0.15
-0.16
0.01
Function 2
2.59
-0.65
0.10
0.41
2.57
0.46
-0.36
-0.71
-0.15
-1.44
2.98
-1.58
0.45
-0.13
0.15
0.04
0.10
-0.10
0.03
0.01
-0.20
-0.32
-0.43
0.43
0.26
0.17
TABLE 4.?Results of stepwise discriminant function analysis on eleven male
anatomical variables in terms of standardized weights and correlation of
variable with discriminant function.
TABLE 6.?Classification results of discriminant function analyses expressed
as per cent of specimens correctly classified.
Variable
CTW
TS/BL
L2
BL
L3/L2
GLR
LFI
WFI
CTL
LPR/BL
L3
Function 1
0.10
-0.04
0.48
0.17
-0.58
0.87
-0.21
-0.46
-0.05
0.56
0.28
0.05
-0.08
0.44
0.03
-0.06
0.68
-0.03
0.05
0.12
0.16
0.28
Function 2
0.21
0.78
-0.88
0.81
-0.85
0.05
-0.49
0.20
-0.31
-0.06
1.24
0.32
0.44
-0.04
0.65
-0.05
0.03
-0.25
0.04
0.37
-0.30
-0.04
TABLE 5.?Results of stepwise discriminant function analysis of thirteen female
anatomical variables in terms of standardized weights and correlation of
variable with discriminant function.
Variable
CTL
BAG
BUW
COP
CGW
FI
AGL
OV
SRL/BUL
OSL
BDW
OCT
OSL/CTL
Function 1
-0.59
-0.26
0.16
-0.28
0.26
0.25
0.49
0.29
-0.07
1.01
0.62
0.56
-1.29
0.49
0.12
0.23
0.15
0.33
0.39
0.40
0.26
-0.01
0.29
0.36
0.38
-0.11
Function 2
0.40
-0.19
-0.70
0.73
0.14
0.19
-0.27
-0.29
0.70
-0.19
0.34
-0.11
0.70
0.14
-0.40
-0.29
0.31
0O3
0.17
0.09
-0.09
0.52
0.16
0.03
0.01
0.08
Data set
Male shell
Female shell
Male anatomy
Female anatomy
Grouped cases Ungrouped cases
89(n = 1
93 (108)
100(60)
100(60)
14 (n = 104)
15(103)
29 (28)
19 (27)
to differences in sample size: more shells per group were
measured than animals (seven to eleven vs. five), which results
in increased variance in discriminant function scores for the
former.
Superior separation of groups based on the first two
discriminant functions for male anatomical data reflects
relative diversity and ease of quantification of penial morphol-
ogy compared to shell and female anatomy. Morphologic
diversity among shells of the congeners, apart from size and
general form, is slight (Figures 3, 11, 16, 21, 31, 36). In
addition, shell variables used, consisting of those traditionally
employed by malacologists, are admittedly unsuited for
accurate quantification of shell form, which is far better
achieved using parameters developed by Raup (1961, 1966).
Female reproductive morphology is similarly conservative,
with little variation in pallial oviduct and bursa copulatrix
complex apart from size (and absence of seminal receptacle in
P. sancarlosensis) (Figures 10, 26). Penial variation is more
pronounced, and involves easily quantified features that
include number of glandular ridges and relative sizes of lobe
and filament (Figures 8, 9, 13, 15, 18, 20, 23, 25, 28, 30, 33,
35, 38).
The relatively high frequency of misclassification of
ungrouped specimens is partly a result of interpopulation
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
variation within species. Significant variation in shell size and
general form is evident in several species (Figures 3d-g, 21g-i,
?>la-g, 36) and even occurs among samples of P. montezu-
mensis from Montezuma Well proper and its upper outflow,
with classification of the latter only 33% for either sex. Similar
levels of anatomical variation were not readily apparent to us
(note only slight variation in penes among six populations of
P. thompsoni, Figure 38), yet misclassification was still
frequent. This reflects occurrence of subtle variation among
conspecific populations involving complex combinations of
variables. An additional explanation is inflated variation within
populations due to small sample sizes (five to eleven).
Regardless of its cause, existence of such variation suggests
that discriminant function equations solely based on data from
topotypes do not consistently correctly classify specimens from
additional populations.
Addendum
A recent (Taylor, 1987) monograph of springsnails of New Mexico and vicinity, published
subsequent to correction of proofs of this paper, includes descriptions of a few of the
species that we name herein and we recognize the following cases of synonymy: Tryonia
gilae Hershler is a junior primary homonym (and junior subjective synonym) of Tryonia
gilae Taylor; and Pyrgulopsis confluentis Hershler, P. sancarlosensis Hershler, and P.
cochisi Hershler are junior subjective synonyms of Fontelicella trivialis Taylor,
Apachecoccus bernardinus Taylor, and Yaquicoccus bernardinus Taylor, respectively.
Discussion of generic allocations provided by Taylor for some of the above is tabled for
a forthcoming paper by one of us (RH).
Appendices
Appendix 1. Shell data for species of Pyrgulopsls. USNM catalog numbers are given for localities. Sample sizes are given in parentheses
(#males, #females). Asterisked localities have significant (two tailed t-test, p<.05) sexual dimorphism in shell height (SH).
Measurements are in nan. X = mean, S - standard deviation.
Species and locality
P. glandulosus
*847205
(n = 8,7)
P. deserta
*847202
(n =- 8,9)
847206
(n = 8)
P. bacchus
84203
(n - 10)
P. conicus
847237
(n = 9,8)
*847201
(n - 7,9)
P. morrisoni
*847231
(n - 9)
847220
(n =? 8,10)
P. montezumensis
847233
(n = 10,8)
*847230
(n - 10,9)
P. solus
*850290
(n - 8,10)
P. simplex
*847236
(n - 9,10)
847222
(n - 10,9)
P. confluentis
847234
(n = 10)
*847229
(n = 10)
P. sancarlosensis
847226
(n - 13)
847217
(n - 10)
847224
(n - 9)
P. cochisi
*847218
(n - 9)
P. thompsoni
~*847238
(n = 10,9)
847221
(n - 8,9)
*847228
(n - 10)
847227
(n - 8,9)
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
X
s
M
2.
0.
SH
24
18
1.51
0.17
1.
0.
2.
0.
2.
0.
2.
0.
1.
0.
2.
0.
2.
0.
2.
0.
1.
0.
2.
0.
2.
0.
2.
0
1,
0
1,
0
1
0
1
0
1
0
2
0
1
0
2
0
1
0
47
13
57
23
36
17
17
21
89
10
29
19
12
28
06
24
57
,08
,08
.12
.18
.19
.81
.29
.62
.12
.90
.19
.34
.17
.50
.06
.41
.09
.67
.24
.97
.08
. 15
.25
.91
.19
F
2.
0.
1.
0.
1.
0.
2.
0.
2.
0.
2.
0.
2.
0.
2.
0.
2.
0.
2.
0.
1.
0.
2.
0.
2.
0,
3.
0
1,
0
2,
0
1
0
1
0
1
0
2
0
2
0
2
0
1
0
56
15
41
13
76
15
54
16
31
14
47
13
08
14
37
18
18
15
41
14
72
,16
,31
.09
.21
.16
.01
.68
.83
.80
.01
.20
.54
.34
.62
.82
.54
.12
.41
.17
.08
.18
.51
.31
.85
.19
SW
M
1.67
0.13
1.28
0.12
1.15
0.06
1.79
0.08
1.54
0.11
1.43
0.14
1.43
0.07
1.64
0.10
1.41
0.20
1.42
0.12
1.17
0.04
1.43
0.11
1.42
0.11
1.80
0.16
1.30
0.09
1.28
0.08
1.06
0.10
1.11
0.04
0.89
0.05
1.68
0.11
1.43
0.06
1.40
0.10
1. 19
0.08
F
1.93
0.08
1.20
0.10
1.36
0.04
1.84
0.07
1.57
0.09
1.65
0.10
1.54
0.11
1.68
0.11
1.51
0.09
1.66
0.06
1.29
0.12
1.55
0.07
1.56
0.11
1.92
0.15
1.42
0.06
1.33
0.11
1.20
0.16
1.20
0.09
1.10
0.09
1.59
0.09
1.43
0.12
1.62
0. 11
1.23
0.05
AH
M
1.13
0.09
0.85
0.06
0.74
0.04
1.23
0.05
1.11
0.07
1.01
0.09
0.92
0.07
1. 11
0.10
0.95
0.14
0.93
0.07
0.78
0.03
1.01
0.07
0.97
0.09
1. 18
0.10
0.83
0.06
0.92
0.07
0.74
0.07
0.75
0.03
0.63
0.03
1.20
0.08
0.97
0.04
0.96
0.09
0.83
0.06
F
1.28
0.10
0.80
0.06
0.87
0.03
1.23
0.05
1.11
0.08
1.11
0.07
0.98
0.08
1.12
0.07
0.99
0.05
1.07
0.06
0.87
0.08
1.08
0.07
1.05
0.11
1.24
0.10
0.93
0.04
0.93
0.06
0.83
0.11
0.82
0.05
0.69
0.05
1.12
0.08
0.97
0.06
1.12
0.08
0.87
0.04
Parameter!
AV
M
1.03
0.10
0.75
0.05
0.71
0.04
1.13
0.05
0.96
0.07
0.84
0.08
0.83
0.05
0.97
0.08
0.87
0.10
0.81
0.07
0.73
0.03
0.87
0.06
0.87
0.007
1.10
0.11
0.77
0.04
0.82
0.05
0.67
0.06
0.73
0.03
0.57
0.03
1.05
0.07
0.87
0.04
0.90
0.07
0.75
0.07
1
F
1.
0.
0.
0.
0.
0.
1.
0.
0.
0.
0.
0.
12
06
74
06
82
05
14
07
96
06
98
05
0.91
0.07
0.
0.
0.
0.
0.
0.
0.
0.
0.
0.
0,
0
1,
0
0,
0
99
06
93
08
95
04
82
,07
,94
.06
,90
.06
.16
.09
.84
.03
0.85
0.04
0,
0
0
0
0
0
1
0
0
0
1
0
0
0
.75
.11
.77
.05
.63
.05
.00
.07
.87
.05
.01
.08
.76
.04
LBW
M
1.84
0.14
1.30
0.11
1.22
0.08
2.02
0.12
1.83
0.12
1.66
0.12
1.49
0.10
1.80
0.14
1.55
0.19
1.54
0.19
1.29
0.05
1.67
0.09
1.65
0.15
2.09
0.18
1.36
0.09
1.56
0.14
1.17
0.12
1.28
0.05
1.11
0.05
2.03
0.13
1.60
0.08
1.64
0.15
1.43
0.12
F
2.13
0.18
1.24
0.12
1.41
0.09
2.02
0.11
1.81
0.10
1.88
0.06
1.63
0.10
1.82
0.12
1.65
0.09
1.74
0.08
1.45
0.14
1.80
0.10
1.68
0.08
2.18
0.19
1.50
0.10
1.66
0.90
1.34
0.21
1.40
0.11
1.21
0.08
1.90
0.12
1.59
0.10
1.89
0.17
1.47
0.07
!?
1.
0.
1.
0.
0.
0.
1.
0.
1.
0.
1.
0.
1.
0.
1.
0.
1.
0.
1.
0,
1,
0.
1,
0.
1,
0
1,
0
1,
0
1,
0
0
0
0
0
0
0
1
0
1
0
1
0
1
0
WBW
1
42
13
05
10
99
04
58
08
29
08
23
10
14
05
39
08
25
14
17
.12
.00
.03
.18
.08
.25
.08
.65
.14
.08
.08
.09
.08
.83
.10
.95
.03
.81
.04
.43
.08
.18
.05
.29
.10
.09
.07
F
1.60
0.10
0.95
0.07
1.16
0.06
1.58
0.06
1.33
0.09
1.37
0.05
1.26
0.08
1.43
0.08
1.32
0.08
1.40
0.09
1.08
0.08
1.27
0.08
1.26
0.06
1.76
0.18
1.21
0.06
1.15
0.07
0.67
0.15
1.02
0.08
0.88
0.05
1.42
0.11
1.18
0.08
1.47
0.10
1.13
0.06
CD
M
0.18
0.05
0.09
0.01
0.13
0.10
0.20
0.03
0.13
0.02
0. 13
0.01
0.11
0.02
0.15
0.02
0.15
0.03
0.11
0.02
0.11
0.02
0.13
0.01
0.13
0.01
0.19
0.03
0.14
0.02
0.15
0.02
0.08
0.01
0.11
0.02
0.11
0.01
0.18
0.02
0.12
0.01
0.14
0.02
0.14
0.04
F
0.19
0.01
0.12
0.03
0.12
0.02
0.18
0.03
0.15
0.02
0.15
0.02
0.12
0.02
0.14
0.01
0.17
0.02
0.15
0.02
0.11
0.02
0.14
0.02
0.13
0.02
0.24
0.07
0.15
0.03
0.15
0.03
0.11
0.02
0.13
0.03
1.12
0.02
0.14
0.02
0.13
0.01
0.15
0.01
0.14
0.02
AB
M
0.68
0.07
0.41
0.04
0.42
0.04
0.78
0.11
0.68
0.07
0.61
0.04
0.56
0.03
0.67
0.07
0.59
0.05
0.56
0.06
0.46
0.04
0.59
0.04
0.63
0.07
0.82
0.09
0.50
0.05
0.56
0.07
0.40
0.05
0.45
0.03
0.43
0.03
0.77
0.06
0.58
0.02
0.62
0.07
0.54
0.05
F
0.78
0.05
0.40
0.05
0.50
0.06
0.74
0.06
0.66
0.04
0.69
0.04
0.59
0.05
0.66
0.06
0.62
0.05
0.65
0.65
0.50
0.05
0.63
0.05
0.67
0.11
0.85
0.14
0.53
0.04
0.62
0.06
0.46
0.11
0.48
0.07
0.48
0.04
0.70
0.05
0.59
0.04
0.73
0.07
0.57
0.05
NW
M
3.78
0.16
3.44
0.18
3.47
0.16
4.18
0.12
4.11
0.13
4.07
0.31
3.78
0.15
4.13
0.19
4.00
0.20
4.05
0.37
3.50
0.13
3.81
0.21
4.03
0.18
4.30
0.20
3.73
0.14
3.90
0.26
3.40
0.21
3.33
0.13
3.56
0.21
4.53
0.18
4.03
0.21
4.05
0.23
3.91
0.38
F
3.89
0.13
3.28
0.20
3.69
0.29
4.15
0.18
4.09
0.13
4.25
0.13
4.00
0.13
4.03
0.22
4.06
0.26
4.14
0.13
3.73
0.18
4.00
0.20
4.00
0.18
4.35
0.32
3.68
0.21
4.06
0.17
3.65
0.27
3.53
0.36
3.69
0.17
4.14
0.13
4.03
0.20
4.35
0.29
3.89
0.18
NUMBER 459 61
Appendix 2. Female anatomical data for species of Pyrgulopsis. USNM catalog numbers are given for the localities. For all samples, n - 5.
Measurements are in mm. X = mean, S = standard deviation.
Parameters
Species and locality BL OV OSL OSW OCT CTL CTW FI CGL CGW AGL AGW COL COP COD BUT BUW BDW BAG SRL 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
847202 X 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
S 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
S 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
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
S 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 X 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
S 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 X 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. montezumensis
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
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
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
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
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
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
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
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
S 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
847233
P. solus
850290
P. simplex
847236
P. confluentis
847234
P. sarcarlosen;
847226
X
s
X
s
X
s
X
s
iis
X
s
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
X 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
X 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
6 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Appendix 3. Male anatomical data for species of Pyrgulopsls. USNM catalog numbers are given for the localities. For all samples, n = 5.
Measurements are in mm. X = mean, S = standard deviation.
f. glandulosus
Parameters
Species and Locality BL TS OSL OSW OCT CTL CTW FI LPR WPR L2 L3 PW LFI WFI GLR
847205 X 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
847202 X 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
P. bacchus _
"847203 X 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
S 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^ . conlcus
"847237 X 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
S 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 X 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
S 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 X 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
S 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_. montezumensis
847233 ~ X 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
S 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
2_. solus
"850290 X 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
847236 X 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
S 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^. confluentis
847234 X 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_. sarcarlosensls
847226 " " X 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 X 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
S 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
847218 X 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. thompsonl
"847238 " X 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
S 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
8 4 7 2 2 8
 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
8 4 7 2 2 7
 X 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
S 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
Literature Cited
Bcquaen, J.C., and W.B. Miller
1973. The Mollusks of the Arid Southwest: With an Arizona Check List.
xvi + 271 pages, 7 figures. Tucson: The University of Arizona
Press.
Cole, G.A., and W.T Barry
1973. Montezuma Well, Arizona, as a Habitat. Journal of the Arizona
Academy of Science, 8:7-13.
Ilcndrickson, D.A., and W.L. Minckley
1985. Cienegas?Vanishing Climax Communities of the American
Southwest. Desert Plants, 6:131-175.
Hcrshler, R.
1985. Systematic Revision of the Hydrobiidae (Gastropoda: Rissoacea)
of the Cuatro Cienegas Basin, Coahuila, Mexico. Malacologia,
26:31-123.
Hcrshler, R., and F.G. Thompson
1987. North American Hydrobiidae (Gastropoda: Rissoacea): Redescrip-
tion and Systematic Relationships of Tryonia Stimpson, 1865, and
Pyrgulopsis Call and Pilsbry, 1886. Nautilus, 101(l):25-32, figures
1-35.
Kohn, A.J., and A.C. Riggs
1975. Morphometry of the Conus Shell. Systematic Zoology, 24:346-359.
Landye, J.J.
1973. Status of Inland Aquatic and Semi-aquatic Mollusks of the American
Southwest. 60 pages. Unpublished report submitted to U.S.
Department of Interior, Bureau of Sport Fisheries and Wildlife,
Office of Rare and Endangered Species, Washington, D.C.
1980. Rediscovery of "Amnicola" deserta Pilsbry (Mollusca: Gastropoda)
and Two New Undescribed Hydrobioid Gastropods from the Virgin
River Drainage, Utah-Arizona [Abstract]. Bulletin of the American
Malacological Union. 1980:70-71.
1981. Current Status of Endangered, Threatened, and/or Rare Mollusks
of New Mexico and Arizona. 35 pages. Unpublished report
submitted to U.S. Department of Interior, Bureau of Sport Fisheries
and Wildlife, Office of Rare and Endangered Species, Albuquerque,
New Mexico.
Mayr, E.
1963. Animal Species and Evolution, xiv + 797 pages. Cambridge: Belknap
Press of Harvard University Press.
Meffe, G.K., and P.C. Marsh
1983. Distribution of Aquatic Macroinvertebrates in Three Sonoran Desert
Springbrooks. Journal of Arid Environments, 6:363-371.
Minckley, W.L
1969. Aquatic Biota of the Sonoita Creek Basin, Santa Cruz County,
* U.S. GOVERNMENT PRINTING OFFICE: 1988-181-717/60013
Arizona. Ecological Studies Leaflet, The Nature Conservancy,
15:1-8.
Pilsbry, H.A.
1916. New Species of Amnicola from New Mexico and Utah. Nautilus,
29:111-112.
Pilsbry, H.A., and J.H. Ferriss
1909. Mollusca of the Southwestern States, El: The Huachuca Mountains,
Arizona. Proceedings of the Academy of Natural Sciences of
Philadelphia, 1909:495-516.
Raup, D.M.
1961. The Geometry of Coiling in Gastropods. Proceedings of the
National Academy of Sciences, 47:602-609.
1966. Geometric Analysis of Shell Coiling: General Problems. Journal of
Paleontology, 40:1178-1190.
Taylor, D.W.
1966. A Remarkable Snail Fauna from Coahuila, Mexico. Veliger,
9:152-228.
1970a. Western Freshwater Mollusks, 4 [Editor's summary]. In, A.H.
Clarke, editor. Proceedings of the American Malacological Union
Symposium on Rare and Endangered Mollusks. Malacologia,
10:33.
1970b. West American Freshwater Mollusca, 1: Bibliography of Pleistocene
and Recent Species. San Diego Society of Natural History, Memoir,
4: 73 pages.
1975. Index and Bibliography of Late Cenozoic Freshwater Mollusca of
Western North America. Museum of Paleontology, University of
Michigan, Papers on Paleontology, 10: 384 pages.
1983. Status Investigation of Mollusks. 81 pages. Unpublished report to
New Mexico Department of Game and Fish.
1987. Fresh-water Molluscs from New Mexico and Vicinity. Bulletin of
the New Mexico Bureau of Mines and Mineral Resources, 116: 50
pages.
United States Department Of Interior (USDI)
1984. Endangered and Threatened Wildlife and Plants; Review of
Invertebrate Wildlife for Listing as Endangered or Threatened
Species. Federal Register, 49:21664-21673.
Wiley, E.O.
1981. Phylogenelics: The Theory and Practice of Phylogenetic Syslema-
tics. xv + 439 pages. New York: John Wiley and Sons.
Williams, J.E., D.B. Bowman, J.E. Brooks, A.A. Echelle, R.J. Edwards,
D.A. Hendrickson, and J.J. Landye
1985. Endangered Aquatic Ecosystems in North American Deserts with a
List of Vanishing Fishes of the Region. Journal of the Arizona-
Nevada Academy of Science, 20:1-62.
63

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION
Manuscripts intended for series publication receive substantive
review (conducted by their originating Smithsonian museums or
offices) and are submitted to the Smithsonian Institution Press
with Form SI-36, which must show the approval of the appropriate
authority designated by the sponsoring organizational unit. Re-
quests for special treatment?use of color, foldouts, case-bound
covers, etc.?require, on the same form, the added approval of
the sponsoring authority.
Review of manuscripts and art by the Press for requirements
of series format and style, completeness and clarity of copy, and
arrangement of all material, as outlined below, will govern, within
the judgment of the Press, acceptance or rejection of manuscripts
and art.
Copy must be prepared on typewriter or word processor,
double-spaced, on one side of standard white bond paper (not
erasable), with 1 VA" margins, submitted as ribbon copy (not
carbon or xerox), in loose sheets (not stapled or bound), and
accompanied by original art. Minimum acceptable length is 30
pages.
Front matter (preceding the text) should include: title page
with only title and author and no other information, abstract page
with author, title, series, etc., following the established format;
table of contents with indents reflecting the hierarchy of heads in
the paper; also, foreword and/or preface, if appropriate.
First page of text should carry the title and author at the top
of the page; second page should have only the author's name
and professional mailing address, to be used as an unnumbered
footnote on the first page of printed text.
Center heads of whatever level should be typed with initial
caps of major words, with extra space above and below the head,
but no other preparation (such as all caps or underline, except for
the underline necessary for generic and specific epithets). Run-in
paragraph heads should use period/dashes or colons as neces-
sary.
Tabulations within text (lists of data, often in parallel columns)
can be typed on the text page where they occur, but they should
not contain rules or numbered table captions.
Formal tables (numbered, with captions, boxheads, stubs,
rules) should be submitted as carefully typed, double-spaced copy
separate from the text; they will be typeset unless otherwise
requested. If camera-copy use is anticipated, do not draw rules
on manuscript copy.
Taxonomic keys in natural history papers should use the
aligned-couplet form for zoology and may use the multi-level
indent form for botany. If cross referencing is required between
key and text, do not include page references within the key, but
number the keyed-out taxa, using the same numbers with their
corresponding heads in the text.
Synonymy in zoology must use the short form (taxon, author,
year: page), with full reference at the end of the paper under
"Literature Cited." For botany, the long form (taxon, author,
abbreviated journal or book title, volume, page, year, with no
reference in "Literature Cited") is optional.
Text-reference system (author, year:page used within the text,
with full citation in "Literature Cited" at the end of the text) must
be used in place of bibliographic footnotes in all Contributions
Series and is strongly recommended in the Studies Series
"(Jones. 1910:122)" or "...Jones (1910:122)." If bibliographic
footnotes are required, use the short form (author, brief title, page)
with the full citation in the bibliography.
Footnotes, when few in number, whether annotative or biblio-
graphic, should be typed on separate sheets and inserted imme-
diately after the text pages on which the references occur. Exten-
sive notes must be gathered together and placed at the end of
the text in a notes section.
Bibliography, depending upon use, is termed "Literature Cited,"
"References," or "Bibliography." Spell out titles of books, articles,
journals, and monographic series. For book and article titles use
sentence-style capitalization according to the rules of the lan-
guage employed (exception: capitalize all major words in English).
For journal and series titles, capitalize the initial word and all
subsequent words except articles, conjunctions, and prepositions.
Transliterate languages that use a non-Roman alphabet according
to the Library of Congress system. Underline (for italics) titles of
journals and series and titles of books that are not part of a series.
Use the parentheses/colon system for volume (number):
pagination: "10(2):5-9." For alignment and arrangement of ele-
ments, follow the format of recent publications in the series for
which the manuscript is intended. Guidelines for preparing bibli-
ography may be secured from Series Section, SI Press.
Legends for illustrations must be submitted at the end of the
manuscript, with as many legends typed, double-spaced, to a
page as convenient.
Illustrations must be submitted as original art (not copies)
accompanying, but separate from, the manuscript. Guidelines for
preparing art may be secured from Series Section, SI Press. All
types of illustrations (photographs, line drawings, maps, etc.) may
be intermixed throughout the printed text. They should be termed
Figures and should be numbered consecutively as they will
appear in the monograph. If several illustrations are treated as
components of a single composite figure, they should be desig-
nated by lowercase italic letters on the illustration; also, in the
legend and in text references the italic letters (underlined in copy)
should be used: "Figure 9b." Illustrations that are intended to
follow the printed text may be termed Plates, and any components
should be similarly lettered and referenced: "Plate 9b." Keys to
any symbols within an illustration should appear on the art rather
than in the legend.
Some points of style: Do not use periods after such abbrevi-
ations as "mm, ft. USNM, NNE." Spell out numbers "one' through
nine" in expository text, but use digits in all other cases if possible.
Use of the metric system of measurement is preferable; where
use of the English system is unavoidable, supply metric equiva-
lents in parentheses. Use the decimal system for precise meas-
urements and relationships, common fractions for approximations.
Use day/month/year sequence for dates: "9 April 1976." For
months in tabular listings or data sections, use three-letter abbre-
viations with no periods: "Jan, Mar, Jun," etc. Omit space between
initials of a personal name: "J.B Jones."
Arrange and paginate sequentially every sheet of manu-
script in the following order: (1) title page. (2) abstract, (3) con-
tents, (4) foreword and/or preface, (5) text, (6) appendixes, (7)
notes section, (8) glossary, (9) bibliography (10) legends, (11)
tables. Index copy may be submitted at page proof stage, but
plans for an index should be indicated when manuscript is sub-
mitted.
i