• TABLE OF CONTENTS
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 Front Cover
 Front Matter
 Table of Contents
 Introduction
 Material and mehtods
 Subfamily holospirinae
 Subgenus holospira martens,...
 Holospira maxwelli pilsbry...
 The holospira rehderi species-...
 The holospira albertoi bartsch...
 The holospira zygoptyx
 The holospira melea species-gr...
 Holospira hyperia Bartsch 1926
 Back Cover






Group Title: Bulletin of the Florida Museum of Natural History
Title: Urocoptid landsnails of the genus Holospira from southern Mexico
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00087419/00001
 Material Information
Title: Urocoptid landsnails of the genus Holospira from southern Mexico
Series Title: Bulletin of the Florida Museum of Natural History ; vol. 45, no. 3
Alternate Title: Urocoptid land snails of the genus Holospira from southern Mexico
Physical Description: p. 63-124 : ill. ; 28 cm.
Language: English
Creator: Thompson, Fred G. ( Fred Gilbert ), 1934-
Mihalcik, Elizabeth L. ( Elizabeth Louise ), 1964-
Publisher: Florida Museum of Natural History
Place of Publication: Gainesville, FL
Publication Date: 2005
Copyright Date: 2005
 Subjects
Subject: Urocoptidae   ( lcsh )
Holospira -- Classification -- Mexico   ( lcsh )
Snails -- Classification -- Mexico   ( lcsh )
Genre: Classification   ( lcsh )
bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Includes bibliographical references (p. 96--97).
General Note: Caption title.
General Note: "Publication date December 15, 2005"- -P. 2 of cover.
Statement of Responsibility: Fred G. Thompson and Elizabeth L. Mihalcik.
 Record Information
Bibliographic ID: UF00087419
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 65173623
issn - 0071-6154 ;

Table of Contents
    Front Cover
        Page i
    Front Matter
        Page ii
    Table of Contents
        Page 63
    Introduction
        Page 64
    Material and mehtods
        Page 65
    Subfamily holospirinae
        Page 66
    Subgenus holospira martens, 1860
        Page 67
    Holospira maxwelli pilsbry 1953
        Page 68
    The holospira rehderi species-group
        Page 69
    The holospira albertoi bartsch 1947
        Page 70
    The holospira zygoptyx
        Page 71
    The holospira melea species-group
        Page 72
    Holospira hyperia Bartsch 1926
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
        Page 78
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    Back Cover
        Page 126
Full Text





FLORIDA
MUSEUM
OF NATURAL HISTORY ,


BULLETIN


UROCOPTID LANDSNAILS OF THE GENUS HOLOSPIRA
FROM SOUTHERN MEXICO



Fred G. Thompson and Elizabeth L. Mihalcik











Vol. 45, \No. 3, pp. 63-124


UNIVERSITY OF FLORIDA


2005


GAINESVILLE







The FLORIDA MUSEUM OF NATURAL HISTORY is Florida's state museum of natural history, dedicated to
understanding, preserving, and interpreting biological diversity and cultural heritage.

The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is a peer-reviewed publication
that publishes the results of original research in zoology, botany, paleontology, and archaeology. Address all inquiries
to the Managing Editor of the Bulletin. Numbers of the Bulletin are published at irregular intervals. Specific volumes
are not necessarily completed in any one year. The end of a volume will be noted at the foot of the first page of the
last issue in that volume.




Richard Franz, Managing Editor
Erika H. Simons, Production



Bulletin Committee
Richard Franz, Chairperson
Ann Cordell
Sarah Fazenbaker
Richard Hulbert
William Marquardt
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Irvy R. Quitmyer
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ISSN: 0071-6154

Publication Date: December 15, 2005






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Managing Editor of the BULLETIN
Florida Museum of Natural History
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PO Box 117800
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Fax: 352-846-0287
e-mail: dfranz@flmnh.ufl.edu








UROCOPTID LANDSNAILS OF THE GENUS HOLOSPIRA
FROM SOUTHERN MEXICO




Fred G. Thompson' and Elizabeth L. Mihalcik2







ABSTRACT

The taxonomy of the landsnail genus Holospira s.s. and subgenus Stalactella from southern Mexico is reviewed. The area of
study includes the state of Puebla and adjacent regions of Morelos, Oaxaca and Veracruz. Stalactella Bartsch, 1906 is redefined
and is recognized as distinct from Propilsbrva Bartsch, 1906 with which it had previously been associated. Twenty-nine species
are recognized. The primary types of all species are illustrated, except for Holospira tetralasma Pilsbry, 1902. New species of
Holospira s.s. include Holospira acanthidia n. sp., H. denserpens n. sp., H. eburnea n. sp., H. zygoptvx n. sp., H. rhinion n. sp.,
H. aurantiaca n. sp., H. scololaema n. sp., H. colymis n. sp., H. fortisculpta n. sp., H. haploplax n. sp. New species of Salactella
include H. cremnobates n. sp., H. psectra n. sp., H. marmorata n. sp., and H. chazumbae n. sp. Holospira goniostoma (Pfeiffer,
1856), H. teres (Menke, 1847), H. hogeana Martens, 1897, H. pfeifferi (Menke, 1847), H. (S.) rose Bartsch, 1906 and Coelostemma
microstoma (Pfeiffer, 1861) are redescribed, and lectotypes are designated. Since its original description the latter species was
referred questionably to Holospira.


Key Words: Urocoptid snails, Holospira, new species, southern Mexico





TABLE OF CONTENTS

Introduction .......................................................... ............... . ................ 64
M material and M ethods .......................................................... .................. 65
Subfam ily H olospirinae .................. ...................................... ................. 66
Genus Holospira Martens, 1860 ...................... .... .................. 66
Subgenus Holospira M artens, 1860 ................................................................... 67
The Holospira goniostoma species-group .......................... ............. 67
Holospira goniostoma (Pfeiffer 1856) ..................................................... ... 67
Holospira nmau well Pilsbry 1953 ................................. ....... .............. 68
The Holospira acanthidia species-group .......................................................68
Holospira acathidia new species ........................................ ...................68
The Holospira rehderi species-group.......................................................... 69
Holospira rehderi Bartsch 1947 ................................. ..... ................ 69
Holospira albertoi Bartsch 1947 .................................... .. ........................ 70
The Holospira zygoptyx species-group ................... ....................71
Holospira zvgoptx new species ........................................ ...................71
The H olospira melea species-group .................................... .................... ... 72
Holospira melea Bartsch 1926 ..................................... ..... ............... 72
Holospira hvperia Bartsch 1926 .................................. ..... ........... ..... 73
Holospira eburnea new species .................................... ..... ............... 73





P.O. Box 117800, Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800 USA fgt(aflmnh.ufl.edu
S2500 Shotwell St., Bainbridge College, Bainbridge, GA 39817, USA emihalcik(lbainbridge.edu

Thompson, F.G., & E.L. Mihalcik. 2005. Urocoptid landsnails of the Genus Holo/pira from southern Mexico. Bull. Florida Mus. Nat. Hist. 45(3):
63-124





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


The Holospira teres species-group ......................... ............................75
Holospira teres (M enke 1847) ................................... .......................... 75
Holospira hogeana Martens 1897 ............................. ...................... 76
Holospira teotitlana Bartsch 1945 ................................. ......................78
The Holospira pfeifferi species-group .................................................... 79
Holospira tetralasma Pilsbry 1902 ..................................... ....................... 79
Holospira pfeifferi (Menke, 1847) ................................. ......................79
Holospira oaxacana Bartsch, 1906 ...................................................... 80
Holospira rhinion new species .................... ...........................81
The Holospira painter species-group .................................. ...................... 82
Holospira painter Bartsch 1906 ........................................... ...................... 82
Holospira denserpens new species ................................ ...................... 82
The Holospira aurantiaca species-group ............................. ............ 83
Holospira aurantiaca new species ............................. ...................... 83
The Holospira scololaema species-group ............................... ........... 85
Holospira scololaema new species .................... ........ .......... ... 85
Holospira colymis new species ................................... .......................... 86
The Holospira fortisculpta species group ............................ ............ 87
Holospirafortisculpta new species ................................................... 87
The Holospira haploplax species-group ........................... ............ 88
Holospira haploplax new species ............................ ........................ 88
Subgenus Stalactella Bartsch 1906 ................................... ........................... 89
Holospira (S.) rosei Bartsch 1906 ....................................... ...................... 90
Holospira (S.) cremnobates new species ............................. ............ 91
Holospira (S.) psectra new species ............................. ..................... 92
Holospira (S.) marmorata new species ............................ ............ 93
Holospira (S.) chazumbae new species ..........................................94
Genus Coelostemma Dall 1895 .......................................................95
Coelostemma microstoma (Pfeiffer 1861) ...........................................95
A cknow ledgm ents ............................................................... ...................... 96
Literature C ited ................................................ ......................................... 96


INTRODUCTION
Holospirinae are an ancient and fascinating group of
organisms found in submesic and xeric habitats from
northern Oaxaca north to southern Texas, New Mexico
and Arizona. Species clearly identifiable as Holospirinae
are recorded from the Paleocene of New Mexico
(Cockerell 1914) and from the Upper Cretaceous of
Alberta (Tozer 1956). Currently 127 species are known
from Mexico, and 26 have been recorded from the United
States. Only a single species, Metastoma roemeri
(Pfeiffer, 1848), occurs in both countries (Florida Mu-
seum of Natural History collection). Holospirids are
confined to calcareous terranes. Most species are obli-
gate inhabitants of limestone or dolomite exposures.
Most have very localized geographic distributions. Some
are confined to areas of a few square meters. Most
others are restricted to areas of a few square kilome-
ters. Usually, specimens are numerous and readily col-
lected when a colony is found. However, populations
may be overlooked easily because very precise ecologi-
cal parameters limit the immediate deployment of colo-
nies. We have found as many as four species occurring
in immediately proximal colonies deployed in different


microhabitats that were separated only by a few meters.
At a locality 1 km east of Azumbilla, Puebla we col-
lected Holospira hogeana Martens, Holospira (s. s.)
sp., Holospira (Stalactella) sp. and Bostrichocentrum sp.
Fifteen genera and subgenera currently are recognized
in the Holospirinae. In most cases their distinctions are
based on shell morphology. Within this subfamily con-
vergence in shell characters appears to have occurred
many times (Thompson 1971, 1988). Distinctions be-
tween subgenera and interpretations of phylogenetic
relationships are rather arbitrary because too little re-
mains known about many of the species comprising the
Holospirinae.
The purpose of the paper is to review two subgen-
era of the landsnail genus Holospira Martens, 1860 from
southern M6xico, Holospira s. s. and Stalactella
Bartsch, 1906. The subgenus Holospira is widely dis-
tributed from Arizona, New Mexico and Texas south to
Oaxaca. Stalactella is restricted to a limited area cen-
tered in Puebla and immediately adjacent Oaxaca. The
geographic scope of this study includes species from
Puebla and the immediate vicinities of Veracruz, Oaxaca





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

and Morelos. Twenty eight species are recognized in
this area, of which fourteen are new. The fauna of
Guerrero does not include any of the species groups we
recognize here, and it will be discussed elsewhere. This
study excludes Bostrichocentrum Strebel, 1880 and most
Coelostemma Dall, 1895 which various authors have
recognized as separate genera or as subgenera of
Holospira.
The needs for this study are two fold. The tax-
onomy of the southern Mexican Holospira s.l. was in a
state of chaos, and a comprehensive review was re-
quired before further progress on the genus can be made
in that part of the country or elsewhere. Many species
from southern Mexico were poorly described, including
some of the earliest names applied to the Holospirinae.
This made it nearly impossible to identify newly collected
material from there. Both Holospira s. s. and
Stalactella had not been defined adequately, and their
relationships to other generic-group taxa remained un-
resolved. We continue to recognize Stalactella as a
subgenus of Holospira. The relationship of Stalactella
to Propilsbrya is less clear. Pilsbry (1953) treats
Stalactella as a section of Propilsbrya. We find that
the similarities of lamellar structure between the two
are overshadowed by numerous other differences in the
shell.
Very few naturalists have collected specimens of
holospirids from the area comprising this study. In gen-
eral this reflects the scant attention that has been given
Mexican landsnails. The names of collectors are unre-
ported for many species described prior to 1900. The
earliest collector recorded to have found holospirids form
southern Mexico was the Prussian naturalist F. Liebmann,
1840-1843. Joseph N. Rose and Joseph H. Painter col-
lected snails during botanical explorations for the United
Sates National Museum between 1903-1905. Charles
R. Orcutt, a botanist from the same institution collected
some mollusks at the same time and later in 1925. Marie
E. Bourgeois, a Mexican naturalist collected mollusks in
the state of Morelos and Puebla between 1939 1947.
Maxwell Smith, a private collector from the United States
collected some mollusks in western Veracruz in 1948.

MATERIAL AND METHODS
This study is based on specimens collected by the au-
thors between 1966 and 2004. Most specimens are de-
posited in the Florida Museum of Natural History. Du-
plicate specimens are deposited in the Instituto Tecnologia
de Ciudad Victoria, Tamaulipas. We have examined the
primary types of all species reported from southern
Mexico, except for Holospira tetralasma Pilsbry, 1902,
the province of which is unknown. All primary types


are illustrated by means of digital photographs repro-
duced at 600 ppi, as are other specimens. Species de-
scribed in the 19'h Century and early 20th Century are
redescribed in cases where earlier descriptions are in-
adequate for identification. Other species described
adequately in the literature are illustrated, and notations
are added where appropriate.
Geographic coordinates. For specimens collected
subsequent to 1990 latitude and longitude were deter-
mined by GPS instrumentation and are given in paren-
theses (). Latitude and longitude for earlier locality
records were taken from gazeteers and are given in
brackets [].
Shell measurements are given in mm and are stan-
dard for elongate-cylindrical shells. The length of the
shell is measured from the tip of the apex to the base of
the peristome. Width is the width of the spire perpen-
dicular to the shell length, but does not include the peris-
tome. The measurements we provide are based on speci-
mens selected to show maximum variation. The follow-
ing abbreviations are used for measurements. L = length
of shell, W = width of shell, AH = aperture height, AW =
aperture width, Wh = number of whorls, RP = ribs on


Figure. 1. Holospira rehderi Bartsch, 1947. An
opened shell showing the positions of the columellar
lamella, parietal lamella, palatal lamella and basal
lamella which comprise the internal barrier.







the penultimate whorl, X = mean, SD = standard devia-
tion, Min = minimum, Max = maximum.
The internal barrier of the shell is viewed by filing
open the lower whorls and selectively chipping away
the shell wall to expose internal structures. In the case
of unique types of species described by 19th Century
authors the barrier was examined by use of radiographs.

Subfamily HOLOSPIRINAE Pilsbry, 1953
Holospirinae are characterized as a group by a com-
bination of traits including the shell, jaw, radula, retrac-
tor muscle system and reproductive anatomy. Holospirid
shells are characterized by having a complete apex at
maturity. The apex remains a functional house for the
digestive gland and gonad. The axis is hollow, although
in some species it may be very narrow and solid in ap-
pearance. The adult shell has an internal barrier, which
usually is confined to the last 1-4 whorls. Basically the
barrier consisting of four lamella: the columellar, the
parietal, the basal and the palatal (Fig. 1). We have
examined numerous species from throughout Mexico.
Modifications through reduction in the number oflamella
occur in the various genera and subgenera. When four
lamellae are present they occur in the combination listed
above. When three only are present the combination
consists of the columellar, parietal and basal lamellae.
When two are present the combination includes the col-
umellar and parietal lamellae. When only a single lamella
is present it is always the columellar lamella. Several
generic groups of Holospirinae lack lamellae. The lamel-
lae develop only at definitive growth of the shell. The
sequence ofontogenetic development precedes first with
(1) the columellar, then (2) the parietal, then (3) the basal,
and last, (4) the palatal. Specimens may have an exter-
nally complete but atypically thin adult shell-form that
has not yet completed development of the internal bar-
rier. Such specimens may lack one or more of the lamella
characteristic of adult shells. The lamellae that are lack-
ing occur in the reverse sequence of their development.
Thus, if one lamella is lacking in a normally 4-lamellate
species, it will be a palatal lamella; if two lamella are
lacking these will be the palatal and the basal, etc. How-
ever, we find that specimens which attain definitive
growth appear to be constant in the number, develop-
ment and location of lamellae characteristic for that taxon.
This is very different from the condition that occurs in
some United States species now placed in the subgenus
Eudistemma Dall, 1895 (see Bequaert & Miller 1973)
in which populations vary in the number of lamella that
may be present (Pilsbry, 1946).
Anatomical information on the Holospirinae is lim-
ited and equivocal. At present it is not possible to re-


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

solve phylogeny and delineate genera on the basis of
available data, and the soft anatomy shows little differ-
entiation among closely related species. This could re-
flect the fact that the species distributions are very re-
stricted because of their obligate confinement to calcar-
eous terrain. Reproductive specialization is not neces-
sary for genetic isolation, nor are other ecological isolat-
ing mechanisms. Table 1 lists the species have been
studied anatomically. We have examined the anatomy
of several of the species discussed below. The results
will be published elsewhere following further study of
Holospirinae from other areas of Mexico.
The Holospirinae may be characterized anatomi-
cally as follow. The jaw is solid and striate. The radula
is generalized in structure and dentition; the transverse
rows of teeth are nearly straight, and the central and
lateral teeth have a single large cusp, the mesocone.
The columellar retractor muscle gives rise lower down
(within two whorls of the mantle collar) to the pharyn-
geal retractor and the right and left ocular retractors.
The ocular retractors each subdivide to form the re-
spective ocular retractor and the pedal retractors; the
right ocular retractor passes through the genital atrium.
The penis retractor muscle originates on the floor of
lung and inserts on the apex of the penis. The penis is
short and bulbous, and it may or may not have a terminal
caecum. A well-developed epiphallus enters the end of
the penis eccentrically. The spermatheca is elliptical
and has a long duct. It is appressed against the uterus at
the base of the albumen gland. The spermatheca duct
is with or without with a caecum.

Genus Holospira Martens, 1860
Holospira Martens; in Albers, 1860: 209. Thompson,
1998: 87-89. International Commission on Zoological
Nomenclature, Opinion 1932: 206-207. (Type spe-
cies: Cylindrella golfussi Menke, 1847, not Cylin-
drella piloceri Pfeiffer, 1841).
Four species of Holospirinae have been described
from the region near Tehuacan, Puebla. These are
Holospira tetralasma Pilsbry, 1902, H. pfeifferi
(Menke, 1847), H. painter Bartsch, 1906 and H.
(Stalactella) rosei Bartsch, 1906. The exact province
of each remains unknown. Holospira tetralasma was
described without locality. The other three species were
described as from Tehuacan. These records can be
interpreted only as approximations, as was frequent with
locality information provided in earlier literature.
Another species, Holospira torrei Pilsbry, 1935,
was described from an unspecified locality in Mexico.
The holotype was collected by a Sefior Botteri, who gave
it to Carlos de La Torre, who in turn gave the specimen





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

Table 1. Species of Holospirinae that have been studied anatomically.


SPECIES


Bostrichocentrum centicostata (Thompson, 1964)
Bostrichocentrum goldmani (Bartsch, 1906)
Bostrichocentrum perplexa (Thompson, 1964)
Bostrichocentrum pupa (Thompson, 1964)
Coelostemma elizabethae (Pilsbry, 1889)
Hendersoniella christmani Thompson & Correa-Sandoval, 1994
Hendersoniella lux lux Thompson & Correa-Sandoval, 1994
Hendersoniella palmer (Dall, 1905)
Holospira (Allocoraphe) minima Martens, 1893
Holospira (Eudistemma) arizonensis Stearns, 1890
Holospira (Eudistemma) chiricahuana Pilsbry, 1905
Holospira (Eudistemma) danielsi Pilsbry & Ferriss, 1915
Holospira (Eudistemma) ferrissi Pilsbry, 1905
Holospira (Eudistemma) nelsoni Pilsbry, 1902
Holospira (Eudistemma) sherbrookei Gilbertson, 1989
Holospira (Eudistemma) tantalus campestris Pilsbry & Ferriss, 1915
Holospira (Holospira) goldfussi (Menke, 1847)
Holospira (Millerspira) miller Gilbertson & Naranjo-Garcia, 1998
Holospira (Millerspira) hoffmani Gilbertson & Naranjo-Garcia, 1998
Holospira (Sonoraloa) remondi laevior Pilsbry, 1953
Holospira (Sonoraloa) dentaxis alamellata Gilbertson, 1993



to Pilsbry. Pilsbry (1935: 2) comments "Sefor Botteri The g'
collected mainly in the state of Vera Cruz, but the local- this group i
ity of this form was not recorded". We have examined eral species
the holotype (ANSP 162325). It belongs to a species- em United
group found in western Coahuila and eastern Chihua- goniostoma
hua. The species is excluded from this study. uncertain.

Subgenus Holospira Martens, 1860 Hol
The following species from Puebla, Veracruz,
Morelos and Oaxaca are assigned to the subgenus Cvlindrella
Holospira. They form twelve or more natural species- 1859: 71
groups. The differences between the groups suggest the Holospira
need for further taxonomic division of the genus. This 1873: 32
must wait until the Holospira in other regions of Mexico figs. 6-6c
are better known. Descr
light tan, ci
The Holospira goniostoma species-group peristome a
The shell is medium sized and tan or light brown in medium siz
color. The apex as well as the spire have distinct axial 0.26 times
ribs that are closely spaced. The spire is slender and moderately
has a moderately elevated, rounded apex. The whorls scalariform
are separated by a shallow suture. The internal barrier arched who
consists of four relatively well-developed lamellae lo- 2.1-2.3 prot
cated in the anti-penultimate and penultimate whorls. and is folio


Thompson, 1964
Thompson, 1906
Thompson, 1964
Thompson, 1964
Pilsbry, 1902
Thompson & Correa-Sandoval, 1994
Thompson & Correa-Sandoval, 1994
Thompson & Correa-Sandoval, 1994
Gilbertson 1989a
Gilbertson, 1989b
Gilbertson, 1989b
Gilbertson, 1993
Gilbertson, 1993
Pilsbry, 1902
Gilbertson, 1989b
Gilbertson, 1993
Pilsbry, 1946
Gilbertson & Naranjo-Garcia, 1998
Gilbertson & Naranjo-Garcia, 1998
Gilbertson, 1993
Gilbertson, 1993


geographic range and species composition of
s not clear. Two species are included. Sev-
from northern Mexico and the southwest-
States have shell features similar to Holospira
.Their relationships to this group remain


ospira goniostoma (Pfeiffer, 1856)
Figs. 2-7
goniostoma Pfeiffer, 1856: 47. Pfeiffer,
0. Pfeiffer, 1862b: 63; pl. 7, figs. 7-9.
goniostoma (Pfeiffer). Fischer and Crosse,
8; pl. 17, fig. 4. Strebel, 1880: 84; pi. 14,
c. Pilsbry, 1902: 77-79; pl. 21, figs. 36-41.
iption.- Shell shiny; color of apex and spire
rossed by lighter tan or white oblique ribs;
nd interior of aperture glossy white. Shell
'ed, 14.1-16.7 mm in length; slender, 0.22-
as wide as high. Whorls, 14.5-17.1. Apex
long and convex with about 9-10 convex or
whorls. Spire cylindrical with moderately
rls. Embryonic shell smooth, consisting of
ruding whorls, the first of which is enlarged
wed by the wider but shorter second whorl.


STUDY







Following whorls of apex crossed by oblique, low, nar-
row ribs that are about a third the width of their
interspaces; ribs with white knobs at their lower ends;
ribs on spire becoming more widely spaced, arched and
less oblique; 39-48 ribs on the penultimate whorl. Um-
bilicus narrowly rimate and partially obscured by the
rounded last quarter of a whorl. Aperture on a short
neck that is about a fifth as long as the width of the last
whorl. Aperture sub-ovate in shape with a weak im-
pression along the parietal wall. Peristome thin but slightly
thickened along its edge, and moderately reflected along
lower palatal, basal and columellar margins. Columella
straight, about a tenth the diameter of the whorls. Inter-
nal barrier consisting of four lamellae, and confined to
the penultimate and anti-penultimate whorls (Figs. 2-5,
radiographs). Columellar lamella one whorl long; low
and rounded; located on the middle of the columella and
confined to the penultimate whorl. Parietal lamella 1.5
whorls long; originating in the lower half of the
penultimate whorl, pendant from the middle of the dor-
sal wall; strongly reflected toward the middle of the outer
wall along its middle. Basal lamella about as thick as
the columellar lamella; occupying the length of the
penultimate whorl; arising vertically from the middle of
the floor; highest near its middle and tapering gradually
at both ends. Palatal lamella consisting of a low ridge
located below the middle of the last half of the
penultimate whorl.
Measurements in mm of the lectotype and the two
paralectotypes are given in Table 2.

Table 2.- Holospira goniostoma (Pfeiffer, 1861)
Measurements of the lectotype and two paralectotypes.
Wh = whorls; Ewh = embryonic whorls; RPW = ribs on
penultimate whorl.


L W AH AW Wh RPW


Lectotype 16.7 3.8 2.3
Paralectotype 15.8 3.5 2.7
Paralectotype 14.1 3.7 -


17.1 45
16.2 39
14.5 48


Type Locality. "Mexico". Lectotype by present
designation: British Museum (Natural History) 1996153.
There are three syntypes in the British Museum. The
largest specimen is designated here in as the Lectotype.
The other two specimens are designated as
Paralectotypes. The province of H. goniostoma re-
mains unknown. We assume the species came from
southern Mexico, but that is not certain.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

Remarks.- This species is distinguished by its me-
dium sized, slender, cylindrical shell with a moderately
long and gently convex apex. The whorls are sculp-
tured with crowded narrow low ribs that are about 1/4-
1/3 as wide as their interspaces. The internal barrier
consisting of four lamella that are confined to penultimate
and anti-penultimate whorls.
Holospira goniostoma is not particularly similar
to other southern Mexican species, although it seems to
be closest to the following species. In size and shape it
resembles the Holospira hogeana species-group (see
below), but those Holospira lack ribs on the spire. Other
species known from southern Mexico are shorter, they
have fewer whorls, and they differ in other particular
characteristics.

Holospira maxwelli Pilsbry, 1953
Figs. 8-10
Holospira maxwelli Pilsbry, 1953: 143; pl. 9, fig. 4
Type Locality.- Veracruz, Beyond Terote
[Perote], near Cordova [Cordoba]. Holotype: ANSP
191105; collected by Maxwell Smith. Two specimens
from the Maxwell Smith Collection are in the Florida
Museum of Natural History (UF 50207) and bear the
datum "beyond Perote".
Distribution.- Known only from the type local-
ity.
Remarks.- Holospira maxwelli tentatively is
placed near H. goniostoma because of the strong simi-
larity in shape and sculpture. Apparently H. maxwelli is
tri-lamellate (Fig. 10). Pilsbry (1953) did not observe a
palatal lamella in the holotype of H. maxwelli, nor did
we find one in the single specimen we opened. Only
three specimens are known for the species, all of which
appear to be mature. It is not certain that the lamella
was very weak and overlooked or inadvertently destroyed
in the two specimens that have been opened.

The Holospira acanthidia species-group
This group consists of a single species. It is char-
acterized by its elongate shape with thin, widely spaced
ribs on the apex and spire. Thee internal barrier con-
sists of three lamella. The parietal lamella bears a single
row of very short denticles.

Holospira acanthidia n. sp.
Figs. 11-16
Description.- Light tan in color with lighter col-
ored ribs which tend to be white on last whorl or two.
Peristome and the interior of aperture white. Shell slen-
der, 11.7-14.1 mm long and 0.22-0.25 times as wide as
high. Whorls 16.1-19.1. Embryonic whorls 2.3-2.5, not





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

offset from subsequent whorls. Apex long attenuated,
consisting of about 10 whorls. Spire with 8-9 whorls,
slightly tapered below. Last whorl only slightly enlarged
with a short descending neck which is about 1/3 1/4 as
long as diameter of preceding whorl (Fig. 16). Neck
with a weak longitudinal impression below periphery.
Umbilicus rimate with a narrow elliptical perforation.
Suture deeply impressed, particularly on the apex. Whorls
of spire weakly arched. Embryonic whorls smooth. Fol-
lowing whorls have strong ribs that are about 1/3-1/4
width of their interspaces. Ribs becoming weaker and
more widely spaced on middle of the spire, but increas-
ing in size on lower two whorls. Penultimate whorl has
20-31 ribs. On neck of last whorl ribs become narrower
and poorly defined on the dorsal and mesad surfaces.
Dorsal side of neck with a weak nearly straight longitu-
dinal impression, causing a weak indentation on parietal
margin within aperture. Aperture broadly auriculate in
shape; about 0.95-1.11 times as wide as high, and about
0.14-0.17 times height of shell. Peristome slightly thick-
ened, narrowly and nearly uniformly reflected around
aperture. Plane of aperture in lateral profile lying at
about 40 to the axis of shell. Columella hollow, 0.12-
0.14 times width of whorls. Internal barrier tri-lamellar,
consisting of the columella lamella, a short basal lamella
and the parietal lamella (Fig. 16). Columella lamella
about 2.0 whorls long; with a thickened periphery, be-
ginning 3/4 of a whorl above aperture and ending in anti-
penultimate whorl where it is most strongly developed.
It extends about 1/4 of distance across the cavity. Oc-
casional small, sharp denticles are present on its sur-
face. Its outer edge deflected upward so that it forms a
shallow gutter along columella. Parietal lamella about
2.5 whorls long beginning in anti-penultimate whorl and
ending at about 3/4 whorl behind aperture; originating
above upper end of columellar lamella by about 1/2 a
whorl. Parietal lamella large, projecting downward about
a third of distance across cavity and then curves abruptly
outward towards palatal wall; bearing a single row of
irregularly spaced, very short, thorn-like denticles on its
crest in posterior half of penultimate whorl. Columellar
lamella about 2.5 whorls long, extending from lower half
of anti-penultimate whorl to upper half of body whorl.
Basal lamella low and thick, about one whorl long; be-
ginning 1.5 whorls above aperture and
Measurements of the holotype and 13 paratypes
(UF 80952) are given in Table 3.
Type Locality.- Puebla, limestone bluff 6.3 km
northwest ofTehuacin; 1750 m altitude. Holotype: UF
34377; collected 9 August, 1965 by Fred G. Thompson;
Paratypes: UF 80952 (17); same data as the holotype.
Snails were collected at the edge of a limestone


bluff on the south side of the road from Tehuacin to
Puebla. The bluff was covered with sparse, bleak, xe-
ric vegetation. Live specimens were found under flat
slabs of limestone.
Remarks.- Like Holospira maxwelli, this spe-
cies differs from H. goniostoma by lacking a palatal
lamella. Holospira acanthidia differs from H.
goniostoma and H. maxwelli by its slender shape and
rugose sculpture. There are about 15-19 whorls. The
apex consisting of about 10 whorls and is slightly con-
vex in outline. The long, slender spire is nearly cylindri-
cal and tends to be constricted below the penultimate
whorl. The sculpture consists of strong, widely spaced
ribs. The tri-lamellate internal barrier also is distinctive.
The long parietal lamella extends through the anti-
penultimate and penultimate whorls. In the anti-
penultimate whorl it projects downward about halfway
across cavity and then flares outward toward palatal
wall. It bears a single row of minute spines. The pari-
etal and columellar lamellae are offset at their upper
ends by about half a whorl. The columella lamella is
short, and the low basal lamella is even shorter.
Etymology.- The species name aKavOa is from
the Classical Greek itSov, a thorn, and the diminutive
suffix idion. The name alludes to the short thorn-like
denticles on the parietal lamella.

The Holospira rehderi species-group
This group of medium-sized species is character-
ized by having thin, translucent shells that are uniform
gray or grayish-brown in color. The apex is attenuated,
and the whorls are sculptured with strong ribs separated
by interspaces that are about 3-4 time as wide as the
ribs. The internal barrier is tetra-lamellate with four
well-developed lamellae that are visible through the shell
wall in fresh specimens. The group is found in extreme
western Puebla, adjacent Morelos and northwestern
Oaxaca.

Holospira rehderi Bartsch, 1947
Figs. 17-39
Holospira rehderi Bartsch, 1947b: 287-288; fig. 2.
Holospira morelosensis Bartsch, 1947b: 288; fig. 4.
Description.- Small to medium in size, 9-15 mm
in length. Shell light brownish-gray in color, thin and
semi-translucent. Typically consisting of 11.5-13.0
whorls separated by a distinctly impressed suture. Sculp-
tured with well-developed slightly oblique ribs through-
out length of apex and spire; ribs vary from crowded
with interspaces scarcely wider than ribs, to having
interspaces 4-5 times width of ribs. Shell shape highly
variable within and between colonies, varying from squat





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Table 3. Holospira acanthidia n. sp. Shell parameters in mm for the holotype (UF 34377) and 13 paratypes (UF


80952). L = length, W = width, AH = aperture height, AW
whorl, X = mean, SD = standard deviation.


aperture width, Wh


whorls, RP = ribs on penultimate


L W AH AW Wh RP W/L AH/L AW/AH


Holotype
Paratypes
X
SD
Min.
Max.


12.8 2.9 2.0 2.0


12.6 3.0 2.0
0.6 0.1 0.1
11.7 2.7 1.8
14.1 3.1 2.1


and barrel-shaped to elongate-cylindrical. Umbilicus
openly rimate. Aperture broadly pyriform-shaped with
a slight node along parietal margin near posterior cor-
ner; aperture projecting forward beyond previous whorl
for about a tenth the diameter of the whorl. Internal
barrier consisting of four lamellae (Figs. 31, 39). Col-
umellar lamella well-developed in penultimate whorl and
extends to first half of last whorl. Parietal lamella be-
gins as a low crest at lower part of anti-penultimate whorl;
in penultimate whorl the lamella extends about halfway
across cavity and curves outward toward outer wall,
and then again it is reduced to a low crest in last half of
penultimate whorl. Basal lamella well developed on
middle of floor of last half of penultimate whorl. Palatal
lamella a low, narrow ridge on lower third of outer wall
in last half of penultimate whorl, and lies opposite gap
between parietal lamella and basal lamella.
Measurements based on the holotypes and three
population sample are given in Table 4.
Type Localities.- Holospira rehderi. Puebla,
Chietla [18 31 'N, 98 34' W]. Holotype: USNM 543589;
collected by Marie E. Bourgeois. USNM 543590 in-
cludes 25 specimens labeled as paratypes.
Holospira morelosensis. Morelos, Tlaquiltenango [18
37' N, 99' 10' W]. Holotype: USNM 543591; collected
by Marie E. Bourgeois. USNM 543592 includes 10
specimens labeled as paratypes.
Distribution. Holospira rehderi is widespread
in Morelos, western Puebla and northwestern Oaxaca
from 1000-2100 m altitude. Morelos: 7.2 km SSW of
Ticumin, 1033 m alt. (UF34285); limestone ridge 2 km
NW of Nopalera (18048'35"N,9903'55"W), 1200 m alt.
(UF 200399); 7 km W ofJojutla, 1033 m alt. (UF 34287).
PUEBLA: 5 km SSE of Izucar de Matamoros,
(1832'34"N, 98 25'40"W); 1340 m alt. (UF 200408,
UF 233192); microwave tower hill 12.5 km SE of Izucar
de Matamoros (18031'16"N, 98024'59"W), 1450 m alt.


18.6 29 0.23 0.16 1.00


17.7
0.8
16.1
19.1


0.24
0.01
0.22
0.25


0.16
0.01
0.14
0.17


1.01
0.05
0.90
1.11


(UF 200081); 12.6 km SSE of Izucar de Matamoros,
1700 m alt. (UF 34382); 13 km. SE of Izucar de
Matamoros, 1475 m alt. (UF 34373). Oaxaca: 2.7 km
W of Teposcolula (17030'22"N, 9729"50"W), 2100 m.
alt. (UF 211612); limestone hill 7 km E of Teposcolula,
km Post 7,2160 m alt. (UF 211630); 2 km NE ofTonald,
1540 m alt. (UF 34361); 13.4 km NE of Tonali, 2060 m
alt. (UF 34380); 13.5 km N of Tonald; 2090 m alt. (UF
34284).
Remarks.- This species occurs in numerous iso-
lated colonies over a relatively large geographic area in
southern Mexico. Such colonies have independent evo-
lutionary histories, and rather distinct morphologically ap-
pearing populations occur throughout this area. Some
populations differ quite strikingly, and the initial impres-
sion is that they represent different species. However,
their morphological characteristics overlap those of other
populations so that no consistent distinctions can be made.
Bartsch (1947b) had before him eleven specimens of
Holospira rehderi and twenty-six specimens of H.
morelosensis. The holotypes are dissimilar in appear-
ance to the extent that Bartsch did not compare the two,
though they were described in the same publication. The
holotype ofH. rehderi is elongate and cylindrical in shape
(Figs. 17-20). It has widely spaced ribs with interspaces
that are 4-5 times as wide as the ribs, and the ribs tend
to be discontinuous across the whorl. The holotype of
H. morelosensis is smaller and is tapered-elongate in
shape (Figs. 21-24). Ithas closely spaced ribs that are
about as wide as their interspaces.
Distinctions between Holospira rehderi and H.
morelosensis are inconsistent within large specimen
samples. The samples presented in Table 3 were se-
lected to show extreme variations between populations.
Other samples bridge the apparent differences between
these. Specimens from 2 km NW ofNopalera, Morelos
(UF 200399, Figs. 25-31) encompass typical variation





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Table 4. Holospira rehderi Bartsch, 1947. Shell measurements of the holotypes of H. rehderi (USNM 543590) and
H. morelosensis (USNM 543591), and parameters of three population samples


W AH AW Wh RP W/L AH/L AW/AH


Holotypes
rehderi
morelosensis


13.6
12.1


13.2 29
12.3 54


0.27 0.18
0.31 0.20


UF 34373 (n = 25). Puebla, 13 km SE of Izucar de Matamoros


X
SD
Min.
Max.


11.62
0.79
9.3
12.2


UF 34287 (n = 12).

X
STD
Min.
Max.

UF 200399 (n = 25).

X
SD
Min.
Max.


W AH AW
3.34 2.08 2.13
0.15 0.29 0.27
3.1 1.6 1.8
3.7 3.2 2.7


Morelos, 7 km W ofJojutla
L W AH AW
13.1 3.5 2.7 2.8
0.89 0.32 0.09 0.11
12.0 2.5 2.7 2.5
14.7 3.8 2.9 3.0

Morelos, 2 km NW ofNopalera
L W AH AW
11.9 3.5 2.4 2.5
0.39 0.12 0.12 0.11
11.3 3.3 2.3 2.4
12.5 3.6 2.7 2.7


for the species. Specimens from 7 km W of Jojutla,
Morelos (UF 34287, Figs. 32-35) have an unusually large
aperture, but a few specimens from the same sample
are typical with regard to this character. Specimens
from 13 km SE of Izucar de Matamoros, Puebla (UF
34373, Figs. 36-39) tend to be short, squat and barrel-
shaped with a larger number of whorls, and have widely
spaced ribs. But variation in this sample includes some
specimens that closely approach in shape and sculpture
the holotype of H. rehderi. We conclude that the ma-
terial we have examined represents a single variable
species, H. rehderi. Holospira morelosensis is a sub-
jectivejunior synonym.

Holospira albertoi Bartsch, 1947
Figs. 40-44
Holospira albertoi Bartsch, 1947a: 141-142; fig. 1.
Description.- Shell light grayish-brown; translu-
cent with the parietal, palatal and basal lamellae visible
through shell in fresh specimens. Medium sized, 12.1-


15.4 mm long; moderately slender, 0.22-0.27 times as
wide as long. Apex attenuated and slightly concave in
outline. Spire cylindrical. Sculpture on post-embryonic
whorls consisting of about 30-76 distinct thin, lamellar
axial ribs. Peristome moderately reflected. Aperture
oblique in lateral profile and extending beyond previous
whorl (Figs. 41, 44). Internal barrier tetra-lamellate.
Measurements of the holotype and 25 specimens
(UF 34383) are given in Table 5.
Type Locality.- Puebla, the steep west bank of a
gully on the SE lower flank of Cerro Chimeco, near
Petlalcingo, very close to the Oaxaca border at km 309
along the Pan-American Highway; 1400-1500 m alt. [18'
04' N, 970 54' W]. Holotype: USNM 543495; collected
by Ing. Alberto R. V. Arellano.
Distribution.- Puebla. Known only from the im-
mediate vicinity of the type locality. Bartsch reports it
from Cerro Sangre de Grado, on the SW side of Km
304 on the Pan-American Hwy. (USNM 543497). We
collected it from a bluff along the Rio Petlalcingo, 1 km


1.06
1.06


Wh
14.46
0.83
12.8
16.1


Wh
12.4
0.41
11.5
13.1


Wh
12.12
0.33
11.9
12.8


RP
37.2
5.18
24
44


RP
45.6
3.50
38
50


RP
42.6
3.04
36
47


W/L
0.32
0.03
0.29
0.36


W/L
0.27
0.03
0.19
0.29


W/L
0.29
0.01
0.28
0.31


AH/L
0.19
0.04
0.17
0.25


AH/L
0.21
0.01
0.19
0.23


AH/L
0.21
0.01
0.20
0.22


AW/AH
1.03
0.13
0.80
1.23


AW/AH
1.02
0.05
0.95
1.12


AW/AH
1.05
0.06
1.00
1.18





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Table 5. Holospira albertoi Bartsch, 1947. Shell parameters of the holotype and of 25 specimens selected to show
variation (UF 34383).


L W AH AW Wh RP W/L AH/L AW/AH


Holotypes
UF 34383
X
SD
Min
Max


15.3 3.7 2.62 2.56 15.4


14.06
0.74
12.1
15.4


3.38
0.13
3.2
3.7


2.63
0.23
2.3
3.3


2.47
0.15
2.1
2.8


SW of Petlalcingo, 1350 malt. (UF 211644, UF 211645),
and at 5 km NW ofPetlalcingo, 1500 m. alt. (UF 34383).
Remarks.- The translucent shell and the attenu-
ate weakly concave apex are distinctive. This species is
similar to Holospira rehderi because of its thin shell, its
medium size, its color and its distinct costate sculpture.

The Holospira zygoptyx species-group
The internal barrier is tetra-lamellate. The basal
lamella unique among holospirids in that it is stout and
flat-topped and has a flange along its outer edge. The
flange projects up and then curves outward to mirror
the curvature of the parietal lamella. The group includes
a single species found near Tehuacan, Puebla.

Holospira zygoptyx n. sp.
Figs. 45-52
Description.- Shell opaque white with a rust col-
ored apex and a white aperture. Medium sized, about
11-14 mm long; relatively slender, 0.24-0.28 times as
wide as long; cylindrical or slightly claviform with a ta-
pered apex that is about a third of the length of the shell.
Whorls 12.9-14.4. Suture moderately impressed. Um-
bilicus rimate or imperforate. Embryonic shell rust col-
ored, smooth consisting of 2.0-2.4 whorls that are nar-
rower but higher the succeeding whorl; not noticeably
offset from following whorls. Apex and spire sculp-
tured with low slightly oblique ribs that are about as wide
as their interspaces; 52-78 ribs on penultimate whorl.
Ribs becoming slightly stronger and more widely spaced
on last whorl. Aperture oblique, lying at an angle of
about 20-25 in lateral profile (Figs. 45, 52); extending
forward slightly beyond body whorl on a short neck.
Dorsal side of neck impressed and nearly smooth. Ap-
erture opening broadly auriculate with a wide embayment
at posterior corner. Peristome moderately expanded
around aperture; narrowest at posterior corner and wid-


15.21
0.62
13.4
16.3


0.24 0.17 0.98


48.72
9.66
30
76


0.24
0.01
0.22
0.27


0.19
0.02
0.16
0.22


0.94
0.08
0.76
1.13


est along columellar margin. Columella narrow, straight,
about 1/10 width of shell. Internal barrier long; consist-
ing of four lamellae confined mostly to anti-penultimate
and penultimate whorls (Figs. 50, 51). Columellar lamella
stout; located on center ofcolumella; extending about 1/
of distance across cavity; confined to penultimate whorl
(Fig 50, the parietal lamella is broken away to show the
columellar lamella). Parietal lamella pendant from middle
of roof; two whorls long, originating at beginning of anti-
penultimate whorl and extending to end of penultimate
whorl, hanging half-way across cavity and then flaring
out toward outer wall along its edge, occluding from view
the columellar lamella (Fig. 51). Basal lamella lying along
middle of floor, confined to lower half of penultimate
whorl; stout, low and flattened with its outer edge raised
as a narrow shelf and curved outward parallel to edge
of the parietal lamella (Figs. 50, 51) Palatal lamella ly-
ing along lower third of middle quarter of penultimate
whorl opposite the gap between the parietal and basal
lamellae.
Measurements of the holotype and 25 paratypes
(UF 337051) are given in Table 6.
Type Locality.- Puebla, 1.0 km by road south of
San Antonia Texcala (18022'42"N, 97027'16"W), 2000
malt. Holotype: UF 190771; collected 10April, 1991 by
Fred G. Thompson. Paratype: UF 337051 (56), ITCV
(25); same data as the holotype.
The type locality lies in an arid scrub-cactus com-
munity on a short and narrow limestone reef with a NE-
SW axis where it is crossed by Highway 125, 15 km
south of Tehuacin on Highway 125 to Huahuapam de
Leon. The limestone reef is bounded on either side by
shales and clays. Snails were found under limestone
boulders at the top of the knoll on the west side of the
road.
Distribution.- Known only from the type locality
Remarks.- This is a tetra-lamellate species with





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

a white opaque shell that is sculptured with slightly ob-
lique axial ribs that are about as wide as their interspaces.
The shell is moderately long and slender with an ex-
tended apex. The aperture is broadly auriculate, and
projects beyond the body whorl for a short distance at
about 25 to the shell axis. The internal barrier is con-
fined to the anti-penultimate and penultimate whorls. This
species is unusual within Holospira by having the pari-
etal lamella pendent downward beyond the columellar
lamella, occluding it from view when the shell is opened.
It is unique within the Holospirinae by having a broad,
flat-topped basal lamella with the outer edge flaring out-
ward upward and toward the outer wall to mirror the
curvature of the parietal lamella.
Superficially Holospira zygoptyx is similar to H.
melea Bartsch, 1926 (see below). The latter is a shorter
and squatter species with poorly defined ribs, the shell is
broadly terete, and it has a tri-lamellate barrier with the
short basal lamella developed as a simple ridge on the
floor of the penultimate whorl.
Etymology.- The species name zygoptvx from
the Classical Greek ~yuS a team or pair, andTirbn, a
fold or leaf, and alludes to the mirrored development of
the parietal and basal lamellae.

The Holospira melea species-group
We group the following three species together in
the Holospira melea species-group because they have
relatively small, opaque, glossy white shells that lack dis-
tinct ribs on the spire. Instead, the sculpture on the spire
consists of low, rounded folds or incremental striations.
The two smooth embryonic whorls are not offset in their
development from the subsequent apical whorls. The
species are tri-lamellate; they lack a palatal lamella. The
parietal lamella is very large, and along its highest sec-
tion its edge curves outward toward the outer wall. The
basal lamella is low and rounded.
Tri-lamellate holospirids occur in scattered regions
of southern Mexico (Puebla, Oaxaca and Veracruz), in
Quer6taro and in Arizona. They have in common the


Table 6. Holospira zygoptyx n. sp. Shell parameters of the


presence of a columellar, a parietal and a basal lamella,
but lack a palatal lamella. They have little else in com-
mon. The tri-lamellate stage appears to have been de-
rived independently in the three geographic regions, and
also it appears to have been independently derived among
some of the species from southern Mexico, such as
Holospira acanthidea and H. maxwelli which are dis-
cussed above. On the basis of the tri-lamellate condition
all of these species could be placed in Malinchea
Bartsch, 1945 (= Tristemma Bartsch, 1906), which
Bequaert and Miller (1973: 141) consider a synonym of
Holospira subgenus Eudistemmma Dall, 1895.
Eudistemma is a group of species found in Arizona and
New Mexico. Species from Puebla appear not to be
closely related to Holospira s. s., and we also doubt
that they are closely related to the subgenus Eudistemma.

Holospira melea Bartsch, 1926
Figs. 53-56
Holospira (Tristemma) melea Bartsch, 1926: 5; pl. 1,
fig. 11. Pilsbry, 1953: 141.
Type Locality.- Puebla, Ixcaquixtla. Holotype:
USNM 363145. Collected by C. R. Orcutt ducting bo-
tanical explorations in southern Mexico.
Distribution.- Known only from the type locality.
The holotype is the only known specimen.
Remarks.- We provide figures of the holotype of
Holospira melea and the following species, Holospira
hyperia Bartsch, 1926. Holospira melea and H.
hyperia are known only from single specimens each.
The holotype ofH. melea is an adult shell in which the
internal barrier appears to have reached definitive de-
velopment. The holotype of H. hyperia may not be
mature, as is suggested by the adnate aperture. Both
species are poorly known. They are readily recognized
by their shell shapes. Measurements for the two holo-
types are as follow.


Length width
Holospira melea 13.9 4.3
Holospira hyperia 13.0 4.6


ApH ApW
2.5 2.7
3.1 2.9


whorls
12.6
14.0


holotype (UF 34373) and 25 paratypes (UF 337051).


L W AH AW Wh RP W/L AH/L AW/AH


14.7 3.6 2.8 2.9


12.44
0.83
11.2
14.1


3.26
0.11
3.0
3.4


2.27
0.29
2.2
3.3


2.35
0.15
2.1
2.6


14.5 63 0.24 0.19 1.04


13.44
0.52
19.9
14.4


66.04
6.52
52
78


0.26
0.02
0.24
0/28


0.18
0.02
0.17
0.20


1.05
0.13
0.98
1.14


Holotype
Paratypes
X
SD
Min.
Max.







Holospira hyperia Bartsch, 1926
Figs. 57-59
Holospira (Tristemma) hyperia Bartsch, 1926: 6; pl. 1,
fig. 15. Pilsbry, 1953: 141.
Type Locality.- Puebla, Esperanza. Holotype:
USNM 363146. Collected by C. R. Orcutt, while con-
ducting botanical explorations in southern Mexico.
Distribution.-Known only from the type locality.

Holospira eburnea n. sp.
Figs, 60-66
Description.- Shell pupiform; cylindrical in shape
with a gently rounded conical apex. Spire glossy white
with a dark-tinged apex in which the embryonic whorls
are reddish-black in live specimens; peristome white;
interior of aperture tinged with orange or light brown.
Shell small-medium in size for a Holospira; up to about
13 mm in length, and 0.28-0.39 times as wide as long
with 10.6-13.5 whorls; spire with 4-6 whorls. Suture
shallow but distinctly impressed; whorls weakly arched
between suture. Embryonic whorls smooth and pro-
truding. Subsequent whorls of apex with oblique, low,
smooth, rounded ribs that are about as wide as their
interspaces. Usually ribs are vague or obsolete on spire,
but in some specimens they may be nearly continuous to
the base. Umbilicus usually imperforate; occasional
specimens rimate. Aperture broadly auriculate in shape
with a weak indentation behind parietal margin near pos-
terior corer; plane of aperture parallel to shell axis;
extending forward slightly beyond previous whorl (Fig.
66). Aperture slightly wider than high and about 0.21-
0.25 times length of shell. Peristome conspicuously re-
flected along outer, basal and columellar margins, nar-
rower along parietal margin. Shell wall thin but strong.
Axis straight-sided and nearly uniformly wide in lower
whorls; 0.12-0.14 times shell width. Internal barrier
confined to penultimate whorl, not visible from within
aperture (Figs. 64, 65). Tri-lamellate; palatal lamella
absent. (One specimen of 14 opened had a very weak


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

elongate deposit where the palatal lamella would be).
Columellar lamella one whorl long; located on center of
axis; thin, blade-like; extending about 1/5 of distance
across cavity. Parietal lamella one whorl long; along its
middle it is pendent from the center of the roof to center
of cavity and then is reflected toward outer wall. Basal
lamella about a half whorl long, located in middle of
penultimate whorl; low and rounded.
Measurements of the holotype and 25 paratypes
(UF 341576) are given in Table 7.
Type Locality.-Puebla, limestone hill-top 7.6 km
south-southwest of Molcaxac (1840.1'N, 97054.2'W);
2040 m alt. Holotype: UF 34294; collected 23 October,
1970 by Fred G. Thompson. Paratypes: UF 341576
(345), ITCV (200); same data as the holotype. UF
200092 (278); same locality, collected 22 February, 1992
by Fred G. Thompson. UF 200589 (70); same locality,
collected 28 October, 1992 by Fred G. Thompson and
Elizabeth L. Raiser.
The type locality is in submesic scrub vegetation
with sparsely scattered small deciduous trees and shrubs.
Holospira eburnea was collected on a limestone hill-
top from under limestone blocks and caliche. Specimens
tended to cluster under stones that lay against small
clumps of vegetation. Snails predominantly were under
stone that lay on limestone bedrock. Less frequently
they were under stone that lay on the lateritic soil.
Holospira eburnea was very abundant on the three
occasions of our visits to the type locality. It is the only
species of snail we found there.
Distribution.- Known only from the type locality.
Remarks.- This is a tri-lamellate species with the
internal barrier confined to the penultimate whorl, and
consists of a low columellar lamella, an enlarged pari-
etal lamella and a low short basal lamella. It is recog-
nized by its small pupiform shell that is glossy white with
a dark apex. The apical sculpture consisting of low,
close oblique ribs that become nearly obsolete on the
spire. The shallow suture separates about 10.5-13.5


Table 7. Holospira eburnea n. sp. Shell parameters of the holotype (UF 34294) and 25 paratypes (UF 341576).


L W AH


AW Wh W/L AH/L AW/AH


12.2 3.9 2.8 2.8 12.0 0.32 0.23 1.00


10.80
1.10
9.4
13.2


3.55
0.18
3.3
3.9


2.36
0.19
2.1
2.9


2.38
0.19
2.1
2.9


11.84
0.80
10.6
13.5


0.33
0.03
0.28
0.39


0.22
0.08
0.21
0.25


1.01
0.06
0.96
1.07


Holotype
Paratypes


SD
Min.
Max.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

whorls. The aperture is broadly auriculate, extending
slightly forward beyond the previous whorl. The peris-
tome is strongly reflected and is separated from the pre-
vious whorl.
Holospira eburnea is similar to H. melea. The
latter is terete in shape with a more elongate apex, and a
deeper suture (Figs. 53-55). The sculpture on the apex
consists of fewer and more widely spaced ribs. The
aperture projects farther forward, and the peristome is
narrower. The internal barrier extends through the anti-
penultimate and penultimate whorls. The parietal lamella
is nearly two whorls in length. On its crest is a single
row of very small thorn-like denticles.
Holospira eburnea is similar to H. hyperia, which
has a stockier shell with an abbreviate apex, the suture
is shallower (Figs. 57, 58) and the peristome is adnate to
the previous whorl. As in H. eburnea, the internal bar-
rier is confined to the penultimate whorl, but the parietal
lamella is much weaker.
Etymology.- The species name eburnea is from
the Latin eburneus, of ivory, alluding to the color of the
shell.

The Holospira teres species-group
This is a group of three large species found in ex-
treme eastern Puebla and the immediate vicinity of
Veracruz. This heterogeneous group is difficult to de-
fine. The species have in common large opaque white
shells with a white apex. The embryonic whorls are rela-
tively large, and grade evenly into the following whorls
without being noticeable offset or protruding. The su-
ture is shallow. The spire is smooth or weakly striate,
although some species may be weakly costate near the
base of the spire. The internal barrier consists of four
lamella in most species. However, the palatal lamella is
highly variable in the degree of its development. In some
forms it is present as a low white callus, and in others it
is obsolete or absent. The parietal lamella is long and it
projects downward half-way across the whorl. Its edge
is reflected outward toward the palatal wall along its
highest section.

Holospira teres (Menke, 1847)
Figs. 67-75
Cylindrella teres Menke, 1847: 1. Philippi, 1847: 5; pl.
3, figs. 5-6. Pfeiffer, 1848: 381-382.
Holospira teres (Menke). Fischer and Crosse, 1873:
327-328. Martens, 1897: 279; pl. 17, fig. 3. Pilsbry,
1902: 104-105; pl. 15, figs. 18-20. Haas, 1933: 271.
Bartsch, 1943: 58.
Holospira teres minor Martens, 1897: 278.
Description.- Shell large and stocky, over 17 mm


long, 0.30 times as wide as high; aperture about 0.20
times height of shell. Lectotype with 13.2 whorls. Apex
rather extended with 2.2 smooth protruding embryonic
whorls, which sit at a slightly oblique angle to shell axis.
Embryonic whorls nearly flattened peripherally with a
weak shoulder; stacked and button-like. Following five
apical whorls have numerous close weakly sigmoid axial
ribs. Whorls of spire smooth and weakly striate with
irregularly spaced incremental striation. Last 1.5 whorls
with numerous close ribs that are higher than wide.
Penultimate whorl with 76 ribs. Umbilicus imperforate.
Aperture free from previous whorl and advanced slightly
on a short neck; slightly wider than high; slightly oblique
to shell axis in lateral profile (Fig. 68). Peristome of
lectotype very thin, and weakly reflected only along pa-
rietal and columellar margins; basal and palatal margins
are thin and chipped to a degree that suggests shell is
not quite mature. Internal barrier consists of three
lamella. Columellar lamella inclined vertically outward
from the center of the columella; low and rounded along
its periphery; 1.5 whorls long with its lower end visible
from within aperture. Parietal lamella thin and pendent
from the center of the parietal wall. Its upper half ex-
tends about half-way across whorl and curves outward
toward palatal wall (Figs. 68, 70). It begins at the same
level as columellar lamella and is one whorl long. Lower
half slightly more than half as high as upper half and it is
vertical to roof of whorl. Basal lamella compressed;
about a half whorl long; slanted slightly toward columella.
It originates forward of origin of parietal lamella and lies
at about a third of the distance from the palatal wall to
the columella. Basal lamella highest opposite high point
of parietal lamella and then becoming a rounded ridge
forward of that point. There is no indication of a palatal
lamella (but see Remarks below).
Measurements in mm of the lectotype (SMF 7023)
and two paralectotypes (BMNH 1996135) are as fol-
low.
length width ApH ApW whorls
Lectotype 17.6 5.2 3.4 3.6 13.2
Paralectotype 20.1 5.7 4.5 4.4 12.6
Paralectotype 22.0 7.8 13.2
Type Locality.- "State ofPuebla". Lectotype by
present designation: SMF 7023; collected by F. Liebmann,
1840-1843. This specimen was cited by Haas (1933,
Archiv Molluskenkunde, 65: 271) as "der Typ". Some
or all of Menke's collection was purchased byA Gysser,
which was subsequently acquired by the Senckenberg-
Museum (R. Janssen, personal communication).
The lectotype is the only specimen in the
Senckenberg-Museum (Figs. 67-70). The Senckenberg-
Museum specimen is selected as the lectotype because







ofHaas's notation. Three syntypes in the British Mu-
seum (Natural History), 1996135, are here in designated
paralectotypes. One is illustrated (Figs. 71-75). They
bear no additional information concerning their prov-
enance. We refer them to Holospira teres, but with
some doubt because of their larger size and more at-
tenuate apex. Radiographs comprising Figs. 72-75 of
the paralectotype illustrated in Fig. 71 show that the in-
ternal barrier is very similar to that of the lectotype.
Distribution.- Unknown.
Remarks.- This species is recognized by its
opaque glossy white color with a white apex. It is large,
stout and cylindrical-conical in shape. The apex is sculp-
tured with numerous close, fine tread-riblets. The spire
is smooth but marked with incremental striation. The
last whorl has coarse axial ribs. The internal barrier is
tri-lamellate and is confined to the penultimate whorl
and the upper half of the body whorl. The columellar
lamella is relatively weak and narrow. The parietal
lamella extends halfway across the cavity and then curv-
ing toward the outer wall. The basal lamella is low and
is about half a whorl long.
This is a large species compared to most other
Holospira known from Puebla. Three aspects of its
shell relate it with other species from the region about
Tecamachalco, Puebla. The shell is glossy white, the
apex is attenuated with protruding embryonic whorls and
the last whorl is sculpture with numerous close, sharp
ribs. The high, outwardly flaring parietal lamella, and
the short but high, thin, basal lamella are similar to
Holospira hogeana Martens, 1897 from near
Tecamachalco. There is no indication of a palatal lamella
in the lectotype, but this may be due to either of two
causes. The shell had been opened previous to when
we examined it, and the palatal lamella may have been
on the portion of the wall that was removed. In this
species-group the palatal lamella is very weakly devel-
oped, and it could easily have been unnoticed. Another
possibility is that the palatal lamella may not yet have
developed in the growth of the shell. The thin lamella of
the lectotype appear to be under-developed. The weak
development of the peristome also indicates that the lec-
totype is slightly immature.
Holospira hogeana Martens, from north of
Tehuacin, near Chapulco and Azumbilla, is a large spe-
cies that is similar in shell-size to H. teres, but the apex
is shorter, the shell is dull white, not glossy as in H. teres,
the palatal lamella is very weak or obsolete, and the
basal lamella when present is a short, low, broadly rounded
ridge. The lectotype ofH. teres lacks a palatal lamella,
but this may be an artifact of various causes. The speci-
mens figured as Cylindrella teres by Pfeiffer (1862a:


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

pl. 6, figs. 28-29) and copied by Pilsbry (1902: pl. 15;
figs. 18-19) appear to be H. hogeana.

Holospira hogeana Martens, 1897
Figs. 76-85
Holospira teres higeana Martens, 1897: 280; pl. 16,
fig. 17. Pilsbry, 1902: 104; pl. 23, fig. 71.
Description of the lectotype (Figs. 76, 81).- This
is an old, weathered, lusterless shell, which in other re-
spects is in good condition. Color uniform white. Mod-
erately large, about 17 mm long; about 0.28 time is wide
as high. Containing about 15 whorls. Spire cylindrical
with a convex apex. Umbilicus narrowly perforate.
Embryonic shell not conspicuously set off from follow-
ing whorls of spire; consisting of 2.5 smooth whorls that
gradually increase in width; first embryonic whorl in-
flated; second whorl shorter but wider than the first.
Apex below embryonic whorls consisting of five whorls
that form an evenly arched dome above spire; apical
whorls strongly arched between deeply impressed su-
ture; sculptured with numerous closely spaced oblique
thread-ribs that are about as wide as their interspaces;
upper ends of ribs recurved posteriorly. Whorls of spire
nearly flat-sided and separated by a weekly impressed
suture. Sculptured as on the apex, except that the ribs
are weaker developed and are more widely spaced until
the last whorl where they become stronger and more
crowded toward the aperture. Aperture extended for-
ward on a very short neck, the length of which is about
0.07 diameter of last whorl. Columella straight, 0.17
times width of shell and concave in outline within each
whorl. Internal barrier consisting of four lamellae. The
following description is based on radiographs of the lec-
totype (Figs. 77-80). Columellar lamella low, rounded,
located on middle ofcolumella; 1.5 whorls long confined
to penultimate whorl and upper half of last whorl. Pari-
etal lamella pendent from middle of the roof in the
penultimate and anti-penultimate whorls; reaching its
greatest height in the upper half of penultimate whorl; it
gradually tapers below, and more abruptly above; along
its highest point it curves outward toward palatal wall.
Basal lamella a low semi-lunar ridge near the periphery
of last half of penultimate whorl. Palatal lamella a low
callused ridge located just below middle of outer wall in
first quarter of penultimate whorl opposite and just be-
low outward curvature of parietal lamella.
Measurements of the lectotype are as follow.
Length, 17.2 mm; width, 4.9 mm.; length of aperture,
3.4 mm; width of aperture, 3.1 mm.; whorls, 15.3; em-
bryonic whorls, 2.5. The fifth whorl is perforated by an
insect hole.
Measurements based on 25 specimens from a lime-





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Table 8. Holospira hogeana Martens, 1897. Measurement based on 25 specimens (UF 190779)..

L W AH AW Wh W/L AH/L AW/AH


16.2 4.9
1.3 0.38
11.9 4.0
18.0 5.8


3.3
0.17
3.0
3.7


3.3
0.39
3.0
4.9


17.3
0.78
15.5
18.2


0.30
0.04
0.25
0.34


0.20
0.02
0.18
0.26


1.20
0.09
0.91
1.12


stone hill 1 km east ofAzumbilla, Puebla (UF 190779)
are given in Table 8.
Live-collected specimens from 1 km northeast of
Azumbilla, Puebla (UF 190779) are described as follow.
Specimens from a near by locality, 1 km NE ofChapulco
are illustrated (Figs. 82-85). Shell uniform white, ex-
cept that the embryonic whorls and the earliest whorls
of spire are light tan. Peristome white. Throat behind
aperture light tan. Large, stocky, about 12-18 mm long;
0.25-0.34 times as wide as high. Cylindrical in shape
with a relatively short convex apex. Embryonic whorls
protruding. Last whorl enlarged with a short neck ex-
tending 1/5 the diameter of shell beyond periphery of
last whorl; rounded below periphery, not pinched inward.
Umbilicus narrowly rimate. Suture shallow. Shell con-
sisting of about 15.5-18.2 whorls. Whorls of spire nearly
flat-sided. Apex consisting of about 8-9 whorls that are
slightly rounded peripherally. Embryonic shell consist-
ing of about 2.5-2.6 smooth whorls. First two embry-
onic whorls strongly protruding. Apical whorls with nu-
merous, fine ribs that are as wide as their interspaces.
These become diminished on first whorl of spire and are
absent or are reduced to weak striations or segments on
middle whorls of spire. Ribs again becoming well de-
fined and stronger on last two whorls, and are distinct
but compressed near peristome. Aperture obovate in
shape with a slight longitudinal indentation in parietal wall
behind posterior corer; parietal lip convex in outline
between columella and posterior corer; aperture 0.28-
0.34 times as wide as high; height of aperture about
0.19-0.23 times length of shell. Peristome narrowly re-
flected around aperture, but not noticeably thickened;
narrowest at posterior corer; widest along basal lip;
plane of aperture in lateral profile parallel vertical. Axis
straight and hollow about 0.18-0.23 times width of whorl,
bearing a low, spiral swelling in its middle. Internal bar-
rier usually tri-lamellate. Columella lamella about 1.5-
2.0 whorls long beginning about 0.6 whorls posterior to
aperture; increasing in size to a low, rounded cord in the
penultimate whorl and gradually diminishing into the anti-
penultimate whorl. Parietal lamella about two whorls
long, beginning one-half whorl behind aperture as a low


ridge on roof of the anti-penultimate whorl and then
rapidly increasing in height and reflected outward to-
wards shell wall as a high narrow blade in penultimate
whorl. Parietal lamella frequently bearing numerous,
very short thorn-like denticles along its crest; denticles
not arranged in any definite pattern. Basal lamella may
bear similar denticles. Basal lamellae very weak and
confined to penultimate whorl where it is usually devel-
oped as a low callus ridge or may be absent. Palatal
lamella almost always absent. When present it consists
of a weak peripheral crest below the middle of the outer
wall of the penultimate whorl.
Type Locality.- Veracruz, Maltrate [18' 48' N,
970 16' W; Alt. 2299 m], on the railway between
Veracruz [City] and the City of Mexico, a little west of
Orizaba, eastern slope of the plateau. Lectotype by
present designation: British Museum (Natural History)
1901.6.22.1903; collector unknown
This lot contains a single specimen. The label with
the specimen states "syntype?". There is no evidence
that Martens had more than a single specimen before
him. In the event that other specimens labeled as
syntypes are discovered the specimen cited above is
herein designated as the lectotype. It conforms very
closely to the figure in von Martens, except that it has
an insect hole in the 5th whorl, which is not indicated in
Martens' figure. His figure has a width/height ratio of
0.284. This is slightly wider than his measurements would
indicate, which give a ratio of 0.25. My measurements
give a ratio of 0.285.
Distribution.- This species occurs in eastern
Puebla from the Sierra de Tecamachalco, near
Tecamachalco, east to the Puebla-Veracruz Escarpment
near Ciudad Mendoza, Veracruz, and is found from 1750-
2430 m altitude. Puebla: 1 km NE of Chapulco, 1940 m
alt. (UF 34376, UF 330927) 1.5 kmNNE Chapulco, 1900
m alt (UF 34292); 3 km NNE Chapulco, 1940 m alt.
(UF 34290); 4 km NNE Chapulco, 1940 m alt. (UF
34291); 5 km NE of San Martin Esperillo (1845'11 "N,
9731'36"W), 2430 m alt (UF 200585); limestone hill 1
km E ofAzumbilla, 23 km N ofTehuacin (18037'23" N,
97023'35" W), 2150 m alt. (UF 190779); 1.5 km WNW


X
SD
Min.
Max.







Azumbilla 2400 m alt. (UF 34288); 3 km NWAzumbilla
(1839'21"N, 9724'37"W), 2100 m alt. (UF 200561);
Cerro San Juanico, 4 km E of Azumbilla (1839'49"N,
9728'19"W), 1800 m alt. (UF 233194); 2 km N of
Tecamachalco, 2300 m alt. (UF 34295); 2 km SE f
Tecamachalco, 2140 m alt. (UF 190764, UF 190753);
3.7 km SE of Tecamachalco, 2300 m alt. (UF 34287);
limestone escarpment 23 km W of the Puebla-Veracruz
state line, 10 km E of Esperanza (ca. 18050'N97025'W),
2350 m alt. (UF 81774). Veracruz: 20 km WSW of
Ciudad Mendoza, 1750 m alt. (UF 34372).
Habitat.- This species occurs in xeric areas with
shrubs, Agave and cactuses, where it is found on soil at
or near the base of limestone slopes. We have collected
it at a number of stations, but we never encountered it
on exposed limestone slopes or ledges. Near Chapulco
it was associated on the soil with Bostrichocentrum sp.
Remarks.- The species is recognized by the fol-
lowing combination of characters. The shell is opaque
white, up to 18 mm long; stocky and bullet shaped with a
relatively short rounded apex. The apex is sculptured
with close distinct rib-threads which become obsolete
on the nearly smooth spire. The internal barrier is tri-
lamellate, and is confined to the anti-penultimate and
penultimate whorls. The parietal and columellar lamel-
lae are well developed. The basal lamella is low or ab-
sent and is confined to penultimate whorl. Generally the
palatal lamella is absent. In occasional specimens it is
indicated by a low callused ridge.
Holospira hogeana is distinguished from H. teres
by its nearly lusterless shell with an abbreviated rounded
apex, and its internal barrier. Generally the basal lamella
is weak, and it is located along the outer periphery of the
floor, not in the center as it is in H. teres.
Holospira hogeana shows considerable variation


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

in shell size from one population to the next. Most speci-
mens closely approximate the measurements given
above. Some population samples lack a basal lamella,
or it is reduced in size to a simple rise on the basal wall.

Holospira teotitlana Bartsch, 1945
Figs. 86-100
Holospira teotitlana teotitlana Bartsch, 1945: 92-93;
fig. 2.
Holospira teotitlana filia Bartsch, 1945: 93-95; fig. 1.
Diagnosis.- Large, up to 17-23 mm in length and
0.24-0.29 times as wide as long. Whorls 12.3-14.1,
weakly arched between suture. Shell solid, opaque;
uniform white or light tan in color; cylindrical in shape
with a moderately short rounded apex. Apex sculptured
with very fine oblique incremental threads or smooth.
Spire generally smooth, but usually bearing distinct ob-
lique axial ribs on last whorl or two. Embryonic whorls
not conspicuously set off from following whorls. Aper-
ture is broad and subcircular with a weakly differenti-
ated posterior corer. Peristome moderately reflected
and projects beyond periphery of last whorl (Figs. 89,
95, 100). Internal barrier tetra-lamellate; confined to
penultimate whorl and first half of body whorl (Fig. 90).
Measurements based on the two holotypes and 25
specimens from UF 34283 are given in Table 9.
Type Localities.- Holospira teotitlana
teotitlana: Oaxaca, foothills ofCerro Blanco (Cerro de
Tizatepec) at Teotitlin del Camino. Holotype: USNM
431954; collected by Maria E. Bourgeois.
Holospira teotitlana filia: Oaxaca, Cerro Tizatepec,
near Ignacio Mejia. Holotype: USNM 431957; collected
by Maria E. Bourgeois. Ignacio Meja (180 06' 02N, 970
07' 41W) is a railroad station ca 5 km SW of Teotitlan
del Camino at 788 m.


Table 9. Holospira teotitlana Bartsch, 1945. Shell parameters of the holotypes of Holospira teotitlana teotitlaa
Bartsch, 1945 (USNM 431954), Holospira teotitlana fila Bartsch 1945 (USNM 431957) and of 25 specimens
selected to show variation (UF 34283).

L W AH AW Wh W/L AH/L AW/AH


Holotypes
teotitlana
filia
UF 34283
X
SD
Min.
Max.


17.0 5.5
19.4 6.0


20.81
0.84
19.0
22.6


5.61
0.20
5.1
6.0


3.6
4.0

4.17
0.33
3.5
4.6


3.7
4.1

4.17
0.36
3.0
4.7


12.5 0.32 0.21 1.03
13.2 0.31 0.21 1.03


13.14
0.45
12.3
14.1


0.27
0.01
0.24
0.29


0.20
0.02
0.16
0.23


1.00
0.06
0.84
1.08





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

Distribution.- This species is found in extreme
eastern Puebla and adjacent Oaxaca in the Tehuacan -
Tomellin Valley, at 790-950 meters altitude. Puebla: 10.7
km N ofTeotilan del Camino; 950 m alt (UF 34383); 2.5
km N ofTeotitlin del Camino (UF 200775); 10 km NW
of Coxcatlin, 880 m alt (UF 337311); 11.5 km SSE of
Coxcatlan (UF 337317). Oaxaca: 2.7 km S, 1.5 km W
of Teotitan del Camino (1806.3'N, 97005.2'W), 810 m
alt (UF 337299); 8 km S ofTeotilan del Camino (1805.2'
N. 9704.6'W), 910 alt. ((UF 200425); 2.5 km N of San
Juan de Los Cues (18003.9'N, 97004.1'W), 850 m alt.
(UF 337284); 1.3 km N of San Juan de Los Cues
(18003.3'N. 9704.4'W) (UF 337305).
Remarks.- Holospira teotitlana is similar to H.
hogeana in the general appearance of the shell. Both
are large species with strong, opaque, uniform whitish
shell. Both have similarly sculptured shells. Both have
non-conspicuously protruding embryonic whorls that are
not offset in the angle of growth nor conspicuously dif-
fering in size from the following whorls of the apex.
This is an unusual condition among southern Mexican
Holospria. Generally, H. teotitlana has a tetra-lamel-
late internal barrier confined to the penultimate whorl
and upper body whorl, whereas H. hogeana has a tri-
lamellate internal barrier that occupies the anti-
penultimate and penultimate whorls.

The Holospira pfeifferi species-group
A group of medium- to large-sized tetra-lamellate
species. The shell is lusterless, light brown or tan and is
sculptured with numerous closely spaced axial riblets.
The group is known from the Tehuacan-Tomellin Valley
in extreme southeastern Puebla and adjacent Oaxaca.
The area around Tomellin, Oaxaca has extensive out-
crops of calcareous strata, which remain virtually unex-
plored for mollusks. The valley is known for its en-
demic Cactacea.

Holospira tetralasma Pilsbry, 1902
Cylindrellapilocerei Pfeiffer. Philippi, 1847: 5; pl. 3,
figs. 7, 8. Pfeiffer, 1848: 382 Pfeiffer, 1859: 710.
Pfeiffer, 1862b: 61; pl. 6, figs. 32, 33. (Not Cylindrel-
la pilocerei Pfeiffer, 1841, a nomen nudum (Opin-
ion 1932: International Commission of Zoological No
menclature. 1999).
Holospira pilocerei (Pfeiffer). Fischer and Crosse,
1873: 329; pl. 17, fig. 5. Strebel, 1880: 82; pl. 5, fig.
1. Martens, 1897: 278. (Not Cylindrella pilocerei
Pfeiffer, 1841).
Holospira tetralasma Pilsbry, 1902: 73-75; pl. 21, figs.
23-27.


Type Locality.- Unknown. Location of type un-
known.
Distribution.- Unknown.
Remarks.- Pilsbry named this species on the ba-
sis of the description and figure given in Strebel. The
location of the specimen upon which the description and
illustration are based is unknown. The characteristics
of size, shape and sculpture as depicted by Strebel sug-
gest that this species is most closely related to Holospira
pfeifferi.

Holospira pfeifferi (Menke, 1847)
Figs. 101-105
Cylindrella pfeifferi Menke, 1847: 1. Philippi, 1851: 6,
figs. 4. Pfeiffer, 1848: 382. Pfeiffer, 1862b: 60; pl.
6, figs. 30-31.
Holospira pfeifferi (Menke). Martens, 1860: 40.- Try-
on, 1868: 313; pl. 15, fig. 34. Fischer and Crosse,
1873: 323-325. (exclusive ofvar. b). Strebel, 1880:
84-85; pl. 13, fig. 12. Martens, 1897: 280. Pilsbry,
1902: 80-81; pl. 22, figs. 49-51.
Description .- Shell bluish gray in color with a
glossy white peristome; interior of aperture faint gray-
ish-brown. Shell terete in shape without marked differ-
entiation between the apex and the spire. Lectotype
with an elongate-conical apex of seven whorls and a
cylindrical spire of five whorls; sides of spire weakly
convex. Moderately large and stocky, about 16.1-17.4
mm long, and about 0.32 times as wide as high. Whorls
12.0-12.7. Suture deeply impressed on apical whorls,
shallow on spire. Apical whorls moderately arched;
whorls on spire weakly arched. Base of last whorl ob-
tusely angulate. Umbilicus rimate. Embryonic whorls
2.5; large, about 1.5 mm in diameter, almost a third the
width of the spire; first embryonic whorl swollen and
button-shaped; second whorl shorter but wider; embry-
onic whorls minutely granular. Postembryonic whorls
uniformly sculptured with strong protracted ribs that are
about as high as wide and are about a third the width of
their interspaces; ribs slightly sigmoid and knobby at their
bases; their upper ends weakly crenulate suture; lecto-
type with 66 ribs on penultimate whorl; paralectotype
with 53 ribs. Aperture nearly circular; flattened across
parietal margin; plane of aperture prosocline, lying at an
angle of 10 to shell axis. Peristome adnate to previous
whorl, and slightly crenulate where it crosses over ribs
on body whorl; peristome uniformly thickened and mod-
erately reflected along outer, basal and columellar lips.
Columella weakly sigmoid, about 0.1 times width of
whorl. Internal barrier consisting of four lamellae. The
following description is based on radiographs of the lec-
totype (Figs. 102-105). Columella lamella confined to







penultimate whorl; thick, about 1/5 width of cavity and
protruding from columella at about a fourth of the dis-
tance above floor; lower end visible from within aper-
ture. Parietal lamella pendent from the middle of roof
and confined to penultimate whorl; gradually tapered
along its upper half; thickened along outer edge, which
is strongly reflected toward the outer wall of the whorl.
Basal lamella projecting as a short narrow blade from
the middle of the floor in last quarter of penultimate whorl
and lying immediately below the parietal lamella. Pala-
tal lamella consisting of a low rounded callus one half a
whorl long, and located just above floor of last half of
penultimate whorl.
Measurements in mm of the lectotype and
paralectotype are as follow.
length width ApH ApW whorls
Lectotype 16.1 5.1 4.0 4.0 12.0
Paralectotype 17.4 5.5 3.9 4.0 12.7
Type Locality.- Puebla, Tehuacan. Lectotype by
present designation: British Museum (Natural History)
1996134. There are three specimens labeled as syntypes
in the British Museum. These are from the Cuming
Collection. The specimen selected as the lectotype most
closely conforms to the description in Menke and most
closely matches the figure in Philippi. The paralectotype
is larger, and its shell is more cylindrical in shape than is
the lectotype. In all other aspects its shell is similar to
the lectotype except that it has a fractured peristome.
The third specimen in the British Museum is a different
species, Holospira teres. Pfeiffer (1862: 60; pl. 6, figs.
30-31) described and figured another specimen which
was in his collection, most of which was destroyed dur-
ing the war of 1939-1944. I have been unable to locate
the specimen. His figure is very similar to the lectotype.
Distribution.- Holospirapfeifferi is known only
from the type specimens. It has not been found since its
original discovery. Tehuacan, as cited in early literature,
referred to the general region and not just to the city.
We have collected at many stations is the area, and we
have not found any population that is identifiable with
this species. However, the similarities of sculpture and
the internal barrier of H. pfeifferi to the following spe-
cies are striking similar.
Remarks.- This species is distinguished by the
following combination of characters. The shell is blu-
ish-gray in color. It is moderately large, 16.1-17.4 mm
long and terete in shape with a well-differentiated apex
and spire. There are 12.0-12.7 whorls sculptured with
strongly costulate; ribs that weakly crenulate the suture
above and are knobby along the base. The embryonic
whorls are large and protruding. The peristome is ad-
nate to the previous whorl. The internal barrier is tetra-


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

lamellate and confined to the penultimate whorl, except
that the columellar lamella extends downward to the point
where it is visible from within the aperture.
Holospira pfeifferi is similar to H. oaxacana
because of its sculpture, shape and internal barrier.

Holospira oaxacana Bartsch, 1906
Figs. 106-109
Holospira oaxacana Bartsch, 1906: 132-133; pl. 4, fig. 5.
Description.- Shell opaque, light tan in color (the
three known specimens are dead-collected shells).
Length up to 18 mm. long. Stout, 0.30-0.33 times as wide
as high. Nearly cylindrical with a moderately extended
convex-conical apex. Whorls 11.0-11.5. Sculpture con-
sisting a numerous close sinuous, oblique axial ribs that
are about as wide as their interspaces; about 100 on
penultimate whorl. Peristome most strongly reflected
along columellar margin; narrow and nearly straight
across parietal margin. Aperture slightly oblique in lat-
eral profile; not projecting beyond periphery of previous
whorl. Columella narrow, nearly straight. Internal bar-
rier tetra-lamellate; confined principally to last half of
penultimate whorl. Palatal lamella weakly developed.
Type Locality.- Oaxaca, Tomellin. Tomellin
[1745'N, 9657'W] is a railroad station near Cuicatlan
at the south end of the Tehuacan-Tomellin Valley. Lec-
totype by present designation: USNM 175085; collected
by C. R. Orcutt, Paralectotypes: USNM 1025749 (2).
Bartsch did not designate a holotype among the
three specimens he had before him. He figured the
largest specimen, which was set aside in a separate vial
accompanied by a label stating "TYPE". We select this
specimen as the lectotype (Fig. 106). The other two
specimens are designated paralectotypes. One
paralectotype is worn nearly smooth (Fig. 107). The
second specimen has fine oblique axial ribs (Figs. 108).
Distribution.- Known only from the type locality.
Remarks.- Bartsch's photograph of the lectotype
is almost featureless. It does not depict the fine, close,
ribbed sculpture of the shell, although he describes it
clearly. Bartsch opened the posterior side of a
paralectotype (Figs. 108, 109) as well as the lectotype
to observe the internal barrier. His description of the
barrier in the lectotype is very accurate, although he did
not illustrate it. The lectotype has four lamellae in the
penultimate whorl. The columellar. parietal, and basal
lamellae are well developed. The palatal lamella is
present as a low ridge. The barrier in the paralectotype
is identical in placement and development to the lecto-
type, except that it lacks a distinct palatal lamella, other
than a slight callus on the internal wall. We figure the
opened paralectotype to illustrate the internal barrier (Fig.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

109). The opening on the posterior side of the shell in
the lectotype is too narrow to permit this feature to be
photographed.

Holospira rhinion n. sp.
Figs. 110-116
Description.- Shell thin but solid, translucent, light
tan, lusterless, rough in appearance, peristome and inte-
rior of aperture white. Large, 13.3-16.9 mm long; stocky,
0.26-0.33 times as wide as long. Terete in shape with
an attenuated apex, widest at top of spire (5-6 whorls
above aperture), and diminishing in width below. Whorls
11.5-14.2; nearly flat sided on spire; separated by a shal-
low suture. About 2.3 embryonic whorls; button-shaped
and protruding, smooth, usually same color as following
whorls. Apex and spire sculptured with fine, generally
oblique riblets that are about as wide as their interspaces;
ribs nearly uniform in development through length of shell
but becoming more strongly developed on last whorl.
Aperture ample and sub-circular with a weak indenta-
tion along parietal margin near posterior corer; aper-
ture usually wider than high; height about 0.19-0.26 times
length of shell; plane of aperture lying at a slight angle to
axis of shell and extending slightly forward beyond pe-
riphery of previous whorl (Fig. 112). Peristome thin;
reflected most noticeably along columellar and parietal
margins; least so along posterior comer and outer lip.
Axis narrow, 0.09-0.11 times width of shell at penultimate
whorl; slowly increasing in size and noticeably twisted.
Internal barrier tetra-lamellate; confined to penultimate
whorl (Figs. 114-116). Columellar lamella thin and hori-
zontal, extending about 1/5 of distance across cavity of
whorl; located on middle of axis in the last half of
penultimate whorl and extended as a weak callus into
the first half of body whorl. Parietal lamella large, a
little more than half a whorl in length, beginning in sec-
ond half of penultimate whorl and extending to begin-
ning of body whorl; suspended from center of roof;


reaching nearly half-way across cavity and then flaring
toward outer wall. Palatal lamella 1/4 of a whorl long,
low but solid; located along the lower suture on the outer
wall so that superficially it resembles a secondary basal
lamella (Figs. 114, 115). Basal lamella confined to sec-
ond half of penultimate whorl; situated on center of basal
wall; high, initially erect and then flaring outward along
its edge.
Measurements based on the holotype and 25
paratypes (UF 342781) are given in Table 10.
Type Locality.- Puebla, a limestone hillside on
the west side of Federal Highway 131, 15 km southeast
of Tehuacan; 1540 m altitude. Holotype: UF 34375;
collected by Fred G. Thompson 8 August, 1965.
Paratypes: UF 342781 (171), UF 342782 (6), ITCV (50);
same data as the holotype.
The type locality was nearly barren of plants, ex-
cept for a few cacti, nettles and occasional mesquite.
Snails were found under loose rock.
Distribution.- Known only from the type locality
and a nearby locality. Puebla: 3 km N of Ajalpan
(1823'05"N, 9718'53"W), 1260 m alt. (UF 200779).
Remarks.- This is a moderately large species with
a rough surface sculptured with numerous fine axial
riblets. The color is tan, lusterless and translucent. The
shell is terete in shape with an attenuated apex. The
aperture is ample and sub-circular in shape. The inter-
nal barrier is etra-lamellate and is confined to the
penultimate whorl and the upper half of the body whorl.
The palatal lamella is short and is located on the outer
wall bordering the lower suture in the penultimate whorl.
Holospira rhinion is unique within the Holospirinae be-
cause of the location of the palatal lamella. Usually the
palatal lamella is low in profile (Fig. 114). Occasionally
the palatal lamella is large to the extent that it nearly
occludes from view the basal lamella when the shell is
opened (Fig. 116).
Etymology.- The name rhinion is from the Clas-


Table 10. Holospira rhinion n. sp. Shell parameters of the holotype (UF 34375) and 25 paratypes (UF 342781


L W AH AW Wh RP W/L


AH/L AW/AH


16.0 4.4 3.2 3.3 14.2 72 0.28 0.20 1.04

14.98 4.28 3.20 3.20 12.77 82.84 0.29 0.21 1.00
0.86 0.10 0.20 0.23 0.60 9.23 0.02 0.02 0.09
13.3 4.1 3.0 2.6 11.6 62 0.26 0.19 0.79
16.9 4.5 3.8 3.9 14.2 103 0.33 0.26 1.26


Holotype
Paratypes
X
SD
Min.
Max.







sical Greek, tviov, a small file or rasp, and alludes to
the over all rough appearance of the shell.

The Holospira painter species-group
This group is found in eastern Puebla in the area
from Tehuacan northwest to Azumbilla, over a distance
of about 25 km. The species are alike in having small,
terete or cylindrical-terete shells with an attenuate apex.
The whorls are separated by a well-impressed suture.
The shell is opaque white or livid white with white em-
bryonic and juvenile whorls, and is smooth or is sculp-
tured with fine, close riblets. The internal barrier is tetra-
lamellate, and is confined principally to the penultimate
whorl and may extend partially into the body whorl.

Holospira painter Bartsch, 1906
Figs. 117-122
Holospira painter Bartsch, 1906: 130-131; pl. 3, fig. 5.
Description.- The smallest known species of
Holospira, 7.6-9.3 mm in length; 0.29-0.36 times as wide
as high. Whorls 9.7-11.2. First embryonic whorl 0.8
mm wide. Shell terete in shape, tapering toward base.
Apex and spire opaque white. Sculptured with fine,
closely spaced oblique riblets that are about as wide as
their interspaces. Columellar lamella well developed,
confined to lower half of penultimate whorl and upper
half of body whorl (Figs. 121, 122). Columellar lamella
and parietal lamella visible from within aperture. Pari-
etal lamella extending through length of penultimate
whorl and upper half of body whorl; large and flaring
outward toward outer wall. Parietal lamella separated
from palatal lamella by a narrow gap in lower half of
penultimate whorl. Palatal lamella is a weak callus on
the middle of the lower half of the penultimate whorl;
visible on outside of shell as a longitudinal narrow white
streak. Basal lamella weak, low and confined to last
quarter of penultimate whorl.
Measurements based on 13 paratypes are given in
Table 11.
Type Locality.- Puebla, Tehuacin. Holotype:
USNM 187675. Paratypes: USNM 1025750 (44), Same
data as the holotype. The type series was collected by


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

Joseph N. Rose and Joseph H. Painter during botanical
explorations between 1903-1905.
Distribution.- This species is known only from
the type locality of"Tehuacan", Puebla.
Remarks.- Holospirapainteri has not been found
since its original discovery.

Holospira denserpens n. sp.
Figs. 123-129
Description.- Shell elongate cylindrical-conical in
shape with an abbreviate-attenuated apex. Small, about
9.4-12.5 mm long; 0.26-0.34 times as wide as long.
Opaque; glossy white with a livid tinge to spire. Apex
and embryonic whorls white. Interior of aperture and
interior of whorls light rusty orange in color. Whorls
10.1-12.5; embryonic whorls relatively large and slightly
protruding; following apical whorls button-shaped and
strongly rounded between sutures; spire consisting of
five strongly arched whorls with a noticeably impressed
suture; whorls becoming increasingly turgid toward the
apex. Shell smooth, sculptured with poorly defined ob-
lique incremental striations that tend to become stronger
on the last whorl; apical whorls devoid of ribs. Umbili-
cus usually imperforate, occasionally rimate. Aperture
height 0.20-0.29 times shell length; broadly auriculate in
shape with a narrow embayment at posterior corner;
noticeably offset laterally at posterior corner; forming a
broad shelf above with an indentation behind the peris-
tome. Aperture extended forward for a short distance
beyond previous whorl (Fig. 129). Plane of aperture
nearly parallel to shell axis. Peristome blunt, weakly
reflected along outer and basal margins; hardly at all
along columellar and parietal margins. Columellar axis
narrow, 0.08-0.10 times width of shell; straight sided.
Internal barrier tetra-lamellate; one whorl long, confined
to last half of penultimate whorl and first half of body
whorl (Figs. 127, 128); not visible from within aperture.
Columellar lamella located one whorl behind aperture;
less than half a whorl in length, strong with a rounded
edge and extending about 1/4 of distance across cavity.
Parietal lamella one whorl long; confined to penultimate
whorl; suspended from middle of roof; extending to near


Table 11. Holospira painter Bartsch, 1906. Shell parameters in mm for 13 paratypes (USNM 1025759).

L W AH AW Wh W/L AH/L AW/AH


X
SD
Min.
Max.


8.15
0.48
7.60
9.30


2.65
0.12
2.50
2.80


1.71
0.09
1.46
1.86


1.84
0.05
1.73
1.86


10.22
0.37
9.7
11.2


0.33
0.02
0.29
0.36


0.21
0.02
0.19
0.23


1.08
0.05
1.00
1.18





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

middle of cavity and flaring outward slightly along its
edge. Palatal lamella very weak, consisting of a low
ridge below middle of outer wall; less than half a whorl
in length and confined to the middle half of the penultimate
whorl. Basal lamella low and rounded; lying directly
beneath the parietal lamella; about halfa whorl in length
between the last quarter of the penultimate whorl and
the first quarter of body whorl.
Measurements based on the holotype and 25
paratypes (UF 341543) are given in Table 12.
Type Locality.- Puebla, a limestone hill 1 km east
of Azumbilla, 23 km north of Tehuacin (1837.2'N,
9723.4"W); 2150 m alt. Holotype: UF 190783; col-
lected 11 April 1991 by Fred G. Thompson. Paratypes:
UF 341543 (78), ITCV (30); same data as the holotype.
Topotypes: UF 190782; same data as the holotype;
specimens preserved in ethanol.
Azumbilla is at the intersection of Hwy. 125 and
the road east to Nicolas Bravo. The type locality is an
isolated conical peak covered with xeric vegetation con-
sisting of cactus, Agave, and shrubs. Snails were col-
lected in the immediate vicinity of a cleft in the west-
facing cliff. Holospira denserpens was found aesti-
vating in small solution crypts in the limestone and under
dried mosses on the limestone cliff. A species of
Bostrichocentrum was common under Agave and ter-
restrial bromeliads in the tallus slope at the base of the
cliff.
Distribution.- Known only from the type locality.
Remarks.- The species is characterized by its
small size, being about 9.5-12.5 m long and relatively
slender with an abbreviate-attenuated apex. It is opaque
with a apex white and a glossy livid-white spire. The
apical whorls are button-shaped and strongly arched
between the suture. The whorls are smooth with ir-
regular, fine, oblique striations. The aperture is broadly
auriculate in shape, offset laterally and has a narrow
posterior comer. The internal barrier is tetra-lamellate
and is confined to penultimate whorls and first quarter


of body whorl. The columellar lamella is stout but very
short. The parietal lamella large and pendant. The palatal
lamella is short and very weak. The basal lamella is
strong, low and rounded.
Holospira denserpens is distinguished from H.
painter by its larger shell, the spire is cylindrical as
opposed to being tapered below, the apex is shorter and
more rounded and the sculpture is smooth in contrast to
the distinct riblets in the latter species. The parietal
lamella does not flare toward the outer wall nearly to
the extent that is does in H. painter, the columellar
lamella is much shorter and the basal lamella is stronger.
Etymology.- The species name denserpens is
taken from the Latin, dens, a tooth, and serpens, a ser-
pent, and alludes to the gleaming white, smooth, slender
shell.

The Holospira aurantiaca species-group
A group relationship of the following new species
is not apparent. It strongly resembles the next two spe-
cies by the glossy white, opaque shell with orange or
reddish upper-most apical whorls, and obsolescent sculp-
ture on the spire. Holospira aurantiaca is smaller than
those two species, it is much more slender, and it lacks
minute bristles lining the interior of the body whorl and
the penultimate whorl.
The relatively small slender size of Holospira
aurantiaca bears some resemblance to H. denserpens.
It differs from the latter species by its orange colored
embryonic and early juvenile whorls, its less rotund
whorls, the presence of narrow but distinct riblets on the
apex, the presence of weakly developed sub-obsolete
ribs on the spire, and the more extensive development
of the internal barrier.

Holospira aurantiaca n. sp.
Figs. 130-136
Description.- Shell opaque white with an orange
or rust-colored apical tip; peristome white; interior of


Table 12. Holospira denserpens n. sp. Shell parameters of the holotype (UF 190783) and 25 paratypes (UF 341543).

L W AH AW Wh W/L AH/L AW/AH


11.8 3.3 2.5 2.4 11.0 0.28 0.21 0.96

10.37 3.12 2.17 2.18 11.33 0.30 0.21 1.01
0.82 0.09 0.14 0.17 0.64 0.02 0.10 0.09
9.4 3.1 2.0 2.0 10.1 0.26 0.20 0.85
12.5 3.3 2.4 2.5 12.5 0.34 0.25 1.22


Holotype
Paratypes


SD
Min.
Max.







aperture and interior of whorls orange tinted. Moder-
ately small-sized; 10.8-13.4 mm long; slender, 0.24-0.31
times as wide as high. Terete, widest point at top of
spire, and gradually tapering toward the base; apex at-
tenuated and slightly convex in outline. Whorls 12.0-
14.6; whorls of spire moderately arched between su-
tures; apical whorls become increasingly turgid toward
apex. Embryonic whorls 2.5, protruding; set at a slight
angle to succeeding whorls; regularly increasing in size;
second embryonic whorl narrower but higher than suc-
ceeding whorl; smooth. Postembryonic whorls of spire
with fine oblique riblets that are about as wide as their
interspaces. Riblets becoming obsolescent on the spire,
but when present are larger and more widely spaced.
Usually axial sculpture becomes better defined as bolder
oblique ribs on penultimate whorl and body whorl. Um-
bilicus rimate or imperforate. Aperture broadly auricu-
late; posterior corner forming a small embayment; aper-
ture offset laterally and forming a rather broad horizon-
tal shelf behind parietal lip; shelf with a conspicuous
central indentation; aperture projecting forward for about
1/5 diameter of previous whorl (Fig. 133); slightly ob-
lique to axis of shell. Peristome thick, blunt, weakly
reflected along the basal, columellar and parietal mar-
gin; becoming very narrow along posterior corer. Axis
straight, narrow, about 0.06-0.08 times width of shell;
expanding noticeably in the anti-penultimate and
penultimate whorls. Internal barrier tetra-lamellate (Fig.
136); not visible from within aperture. Columellar lamella
lying on middle ofcolumella, moderately thin, extending
horizontally for about 1/4 of distance to outer wall; one
whorl in length and confined to penultimate whorl. Pa-
rietal lamella two whorls in length, beginning in anti-
penultimate whorl and extending to upper half of body
whorl; suspended vertically from outer third of roof for
about half of cavity width. Palatal lamella about V/2 a
turn in length in the first half of penultimate whorl; weak;
located along lower third of whorl. Basal lamella one


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

whorl in length, confined to penultimate whorl, located
on center of floor, vertical and about 1/4 height of whorl.
Measurements of the holotype and 26 paratypes
are given in Table 13. Specimens from 2 km north of
Tecamachalco are slightly smaller than those of the type
series, but they are typical in all other aspects of the
shell.
Type Locality.- Puebla, limestone canyon 2 km
southeast ofTecamachalco (1851.4'N, 97042.0'W), 2140
malt. Holotype: UF 190756; collected 9 April, 1991 by
Fred G. Thompson. Paratypes: UF 341545 (134) ITCV
(50); same data as the holotype.
The type locality is in a small canyon on the north-
west end of a limestone mountain that faces southwest.
It is reached by driving for about 2 km up a rough graded
road from the southeast edge of Tecamachalco to a
quarry. The area is arid. It is vegetated with cactus-
Agave-Yucca and some shrubs. Holospira aurantiaca
was very abundant on limestone cliffs under flags of
limestone and under leaves of cliff-dwelling Agave.
Distribution.- Known only from the immediate
vicinity of the type locality. Puebla: 2 km N of
Tecamachalco, 2300 m alt. (UF 34282).
Remarks.- The species is distinguished by its slen-
der shell that is weakly terete with an attenuated apex.
It is moderately small, up to about 13.4 mm in length.
The shell is opaque white, with an orange or rust-col-
ored upper-most apex. The sculpture consisting of fine,
close riblets on the apical whorls. The become stronger
but obsolescent on spire, and then becoming stronger
and more clearly developed on the last two whorls. The
internal barrier is tetra-lamellate and extends from the
anti-penultimate whorl to the body whorl. The parietal
lamella extends through the length of last half of the
anti-penultimate to the first half of the body whorl. The
columellar lamella the basal lamella and the palatal lamella
are confined to the penultimate whorl.
Etymology. The name aurantiaca is from the


Table 13. Holospira aurantiaca n. sp. Shell parameters of the holotype (UF 190756) and 26 paratypes (UF
341545).


L W AH AW Wh W/L AH/L AW/AH

13.2 3.3 2.3 2.3 13.9 0.25 0.17 1.00


Holotype
Paratypes


SD
Min.
Max.


12.01
0.68
10.8
13.4


3.27
0.12
3.0
3.5


2.24
0.15
1.9
2.5


2.21
0.22
1.8
2.5


12.93
0.44
12.0
14.6


0.27
0.02
0.24
0.31


0.19
0.01
0.16
0.21


0.98
0.12
0.84
1.20




THOMPSON and MAHALCIK: Southern Mexican holospirid snails

New Latin, meaning orange colored, in reference to the
color of the upper-most apical whorls, and the interior of
the aperture.

The Holospira scololaema species-group
This is a small group of species found in eastern
Puebla where they occur in isolated colonies. The spe-
cies have opaque, glossy white shells that have obsolete
ribs on the spire. They are characterized as a group by
possessing minute spine-like bristles lining the inside of
the lower-most whorls. The bristles point downward
causing an obstruction of the opening into the shell. Un-
doubtedly this is a barrier evolved to impede the en-
trance of arthropods.

Holospira scololaema n. sp.
Figs. 137-144
Description.- Shell glossy, opaque white on spire
with tan embryonic whorls and early whorls of apex;
peristome white, interior of aperture light tan. Moder-
ately large, 15.4-18.2 mm long, moderately stocky, about
0.25-0.32 times as wide as high. Cylindrical in shape
with an extended convex apex. Last whorl extending
forward on a short neck just slightly beyond body whorl
(Fig. 141); rounded below and flattened above with a
bluntly rounded posterior corner. Umbilicus rimate.
Suture slightly impressed. Shell with 12.0-13.7 whorls.
Spire with 5-6 nearly flat whorls. Apex consisting of 6-
7 whorls. Embryonic whorls 1.5-2.0, smooth, enlarged
and button-like, protruding and higher than succeeding
whorl causing the tip of the apex to be slightly concave
in outline. Succeeding post-embryonic whorls of apex
sculptured with numerous closely spaced oblique ribs
that are about as wide as their interspaces; ribs become
weak and poorly define on the spire, but on last whorl
ribs become well defined and are enlarged behind peris-
tome. Aperture broadly auriculate in shape with a weakly
impressed parietal margin; 0.88-1.11 times as wide as



Table 14. Holospira scololaema n. sp. Shell parameters in
335731).


high; 0.17-0.25 times length of shell. Peristome nar-
rowest at posterior corner and widening along basal and
columellar margins. Axis hollow, slightly twisted with a
slight bulge along its middle; narrow, about 0.09-0.10
times width of shell. Internal barrier bi-lamellate, con-
sisting of a well developed columellar lamella and a weak
or obsolete parietal lamella. Columellar lamella begin-
ning in the penultimate whorl and ending in the last whorl
half a turn posterior to the peristome (Figs. 142. 143);
lower end visible within aperture. Parietal lamella obso-
lete or weakly developed in penultimate whorl. Palatal
and basal lamellae absent. Floor and roof of body whorl
and penultimate whorl lined with forward pointing minute
bristles.
Measurements of the holotype and 24 paratypes
(UF 335731) are given in Table 14.
Type Locality.- Puebla, a limestone mountain side
1 km northeast of Yehualtepec; 2340 m alt. Holotype:
UF 34289; collected 22 October, 1970 by Fred G. Th-
ompson. Paratypes: UF 335731 (135), UF 335727
(SEM), ITCV (25); same data as the holotype.
Snails were found on a steep limestone slope un-
der blocks of limestone and caliche. The surrounding
vegetation consisted or low xeric shrubs dominated with
cactuses and Agave. Other snails found associated with
Holospira scololaema included Euglandina sp. and
Salasiella sp.
Distribution.- Known from the type locality.
Remarks.- The species is recognized by its mod-
erately large sized shell that is relatively slender with an
extended apex and is glossy white. On the spire the flat
sided whorls are nearly uniformly high and wide. The
apical whorls are sculptured with numerous close rounded
ribs, which become poorly defined on most of the spire
but are well developed on the last whorl. Thee aperture
is about as wide as high and lacks a conspicuous
embayment at the posterior corner. The internal barrier
is bi-lamellate with a low columellar lamella low extend-



mm for the holotype (UF 34289) and 24 paratypes (UF


L W AH AW Wh W/L AH/L AW/AH

17.0 4.8 3.4 3.6 13.1 0.28 0.20 1.06


Holotype
Paratypes


SD
Min.
Max.


16.7
0.88
15.4
18.2


4.6 3.4
0.22 0.27
4.3 2.7
5.0 3.9


3.3
0.28
2.8
3.7


13.1
0.50
12.0
13.7


0.28
0.02
0.25
0.32


0.20
0.02
0.17
0.25


0.96
0.07
0.88
1.11







ing from the penultimate whorl to half a whorl behind
the peristome. The parietal lamella is obsolete or weakly
developed in the penultimate whorl. The floor and roof
of the body whorl and penultimate whorl are lined with
minute bristles. Holospira scololaema differs from the
next species by the numerous features that are outlined
in the following diagnosis.
Etymology.- The name scololaema is from the
Classical Greek ocoAog, thorn, and laegoo, throat, al-
luding to the thorny spines lining the roof of the lower
most whorls.

Holospira colymis n. sp.
Figs. 145-150
Description.- Shell glossy, opaque white with light
yellow-brown embryonic whorls; peristome white, inte-
rior of aperture light rust-color. Medium sized, 15.0-
17.6 mm in length. Weakly clavate in shape with a mod-
erately extended apex that is weakly convex in outline.
Top whorl of spire widest and highest; lower whorls
decreasing slightly in width and height until the
penultimate whorl. Shell containing 12.7-14.2 moder-
ately arched whorls with a strongly impressed suture.
Apex consisting of about 5-6 whorls. Embryonic whorls
smooth, protruding and enlarged compared to following
post-embryonic whorl. Next 3-4 whorls of apex sculp-
ture with weakly developed oblique riblets that become
obsolete at top of spire, which is the widest whorl. Riblets
most evident along the upper suture of following whorls
until the antepenultimate or penultimate whorl where they
develop into well defined ribs that continue to the peris-
tome. Base of last whorl bluntly angular-rounded; rimate.
Aperture 0.21-0.24 times length of shell; weakly advanced
beyond previous whorl (Fig. 150). Aperture relatively
narrower, 0.77-0.89 times as wide as high; strongly au-
riculate in shape with a conspicuous embayment at pos-
terior comer. Peristome nearly uniformly narrow; slightly
wider along basal margin; nearly straight across parietal
margin to inner edge of embayment, and then curved
around posterior corner. Axis narrow and straight; about
0.1 times the width of the shell. Internal barrier tetra-
lamellate, consisting of four lamellae in last three whorls,
and short bristles lining roof and floor of last 1.5 whorls
(Fig. 148). Columellar lamella beginning as a low fold in
anti-penultimate whorl. In penultimate whorl it becomes
very much enlarged, and extends about a quarter of dis-
tance across cavity, and then diminishes again to a low
fold in body whorl. Lower end of columellar lamella
visible within aperture from an oblique view. The pari-
etal lamella begins as a low fold in the center of roof 2.5
whorls above aperture. In the last half of the penultimate
whorl the parietal lamella abruptly enlarges, extending


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

down nearly half-way across the cavity and then curv-
ing outward toward the outer wall. Then the parietal
lamella is reduced to a low rounded fold in the following
half whorl. Basal lamella confined to first half of the
penultimate whorl; located on center of floor as a low
but strong fold. Palatal lamella lying along lower third
of outer wall in first half of penultimate whorl as a low
fold apposite gap between parietal lamella and basal
lamella, and opposite columellar lamella. Minute bristles
line roof and floor of last 1.5 whorls above aperture.
Bristles also occur on lower ends of parietal and basal
lamellae.
Measurements of the holotype and the 13 paratypes
are given in Table 15.
Type Locality.- Puebla, a limestone hillside 13.5
km west southwest of San Bartola Teontepec
(1826'03"N, 9736'42"W), 2120 m alt., collected 21 Feb-
ruary, 1992. Holotype: UF 200782; collected 21 Febru-
ary, 1962 by Fred G. Thompson. Paratypes: UF 335728
(9), ITCV (5); same data as the holotype.
Distribution.- Known only from the type locality
and a nearby locality. Puebla: the north facing slope of
a limestone hill about 7 km by road N of Los Reyes
Metzontla (1815'46"N, 9730'22"W); 1800 m alt. (UF
233193).
Habitat.- The type locality was goat-grazed and
covered with sparse xeric scrub vegetation. Snails were
found under caliche slabs and limestone blocks. There
the species was collected well into the dry season. Near
Metzontla snails were found near the end of the wet
season aestivating in small solution crypts on the sur-
face of limestone cliffs and boulders. The surrounding
vegetation consisted of xeric thorn shrubs and Agave.
Remarks.- This is another species with an opaque
glossy white shell. The interior of the aperture is light
rust-colored. The shell is weakly clavate in shape, and
the whorls decrease in width and height below the wid-
est part of spire. The apex is moderately extended, but
less so than in Holospira scololaema. The apical sculp-
ture consists of weakly defines oblique riblets that are
nearly obsolete on the spire until the antepenultimate
whorl where they become enlarged into well developed
ribs. The aperture is relatively narrow and strongly au-
riculate in shape with the posterior corner extended as a
small embayment. The tetra-lamellate internal barriers
located in last three whorls. Short bristles line the roof
and floor of the last 1.5 whorls.
The shot bristles lining the roof and floor within the
last 1.5 whorls suggest a relationship with Holospira
scololaema, but the strongly developed internal barrier
consisting of four lamellae in H. colymis contrasts
strongly with the former species. The two species are





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Table 15. Holospira colymis n. sp. Measurements of the holotype (UF 200782) and 13 paratypes (UF 335728).


L W AH AW Wh


W/L AH/L AW/AH


3.7 3.0 13.6 0.29 0.23 0.81


3.58
0.19
3.4
4.0


3.08
0.12
2.9
3.3


13.45
0.52
12.7
14.2


0.29
0.01
0.27
0.31


0.22
0.01
0.21
0.24


0.86
0.04
0.77
0.89


very similar in general appearance, and are closely af-
filiated geographically.
Etymology.- The name colymis is from the Clas-
sical Greek ioA,6tg meaning an obstruction and re-
fers to the internal barrier.

The Holospira fortisculpta species-group
This group is distinguished by its robust, cylindri-
cal-conical shape with an abbreviated dome-shaped
apex. The shell is solid, opaque white, and is elegantly
sculptured with strong, widely spaced ribs. The internal
barrier is tri- or tetra-lamellate, and is confined to the
penultimate whorl and the upper half of the body whorl.
The single known species occurs at higher elevations in
extreme eastern Puebla.

Holospira fortisculpta n. sp.
Figs. 151-158
Description.- Shell pupiform with an abbreviated
apex and protruding embryonic whorls. Spire cylindri-
cal, consisting of about 6-8 whorls. Apex short, dome-
shaped and strongly convex in outline. Shell medium-
sized, about 11-14 mm long. Robust, about 0.28-0.36
times as wide as high. Shell solid, glossy, opaque, livid-
white, with tan upper apical whorls, a white peristome
and a weakly rust-colored throat deep within the aper-
ture. Whorls 13.3-16.3; weakly arched with a decid-
edly impressed suture. Embryonic whorls 2.2-2.3;
smooth. Subsequent whorls sculptured with prominent
protracted ribs that are about half as wide as their
interspaces; ribs becoming stronger and more widely
spaced below; ribs slightly arched on spire with
interspaces about four times width of ribs; 23-44 ribs on
penultimate whorl. Umbilicus rimate or imperforate.
Aperture ample with a prominently reflected peristome;
very broadly auriculate with a weak embayment at pos-
terior corner; plane of aperture slightly oblique to shell
axis, not protruding beyond preceding whorl (Fig. 158).
Peristome solute, reflected; nearly uniformly wide along


outer, basal and columellar margin; becoming narrower
along parietal margin, which is only slightly separated
from previous whorl. Axis straight and gradually in-
creasing in size; about 0.15-0.20 times width of anti-
penultimate whorl; axis slightly swollen below middle in
anti-penultimate and penultimate whorls. Internal bar-
rier tri- or tetra-lamellate; confined to penultimate whorl
and first half of body whorl (Figs. 155-157); not visible
from within aperture. Columellar lamella one whorl long;
confined to last half of penultimate whorl and first half
of body whorl; lying along lower third of axis, moder-
ately stout, horizontal, low, projecting about 1/4 the width
of cavity. Parietal lamella one whorl long, confined to
penultimate whorl; large; originating vertically from center
of roof and flaring outward slightly along edge. Palatal
lamella generally absent (present in one of six speci-
mens examined); confined to first quarter of penultimate
whorl; weak, apparent as a slight calcareous ridge along
lower 1/4 of outer wall. Basal lamella about a quarter
whorl long, confined to last fourth of penultimate whorl
very close to outer wall; low but stout.
Measurements for the holotype and 25 paratypes
(UF 342779) are given in Table 16.
Type Locality.- Puebla, 5 km northwest ofAtenco,
2650 m alt. Holotype: UF 34293; collected 21 October,
1970 by Fred G. Thompson. Paratypes: UF 342779
(165), ITCV (25); same data as the holotype.
Atenco is a small village located at 190 04.6' N,
970 31.6'W, and at 2500 m alt. The type locality lies
northwest ofAtenco on the road to San Salvador el Seco.
Snails were collected on a hillside from under chips and
blocks of limestone just below a pine forest.
Distribution.- Known only from the type locality.
Remarks. This species is identified by its ro-
bust, medium-sized, pupiform shell with an abbreviated
apex. It is solid, opaque white and glossy, and is sculp-
tured with strong, widely spaced ribs. The aperture is
very broadly auriculate with a widely reflected peris-
tome that is narrowly separated from the body whorl.


Holotype
Paratypes
X
SD
Min.
Max.


16.3 4.7


16.17
1.03
15.0
17.6


4.69
0.13
4.5
4.9





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Table 16. Holospirafortisculpta n. sp. Shell parameters of the holotype (UF 34293) and 25 paratypes (UF
342779).

L W AH AW Wh RPW W/L AH/L AW/AH


Holotype
Paratypes
X
SD
Min.
Max.


12.2 4.1 2.8 2.8 14.9 39 0.34 0.23 1.00


12.37
0.73
11.2
13.9


3.99
0.15
3.8
4.3


2.59
0.17
2.1
2.8


2.76
0.18
2.3
3.2


14.81
0.71
13.3
16.3


33.2
6.57
23
44


0.32
0.02
0.28
0.36


0.22
0.04
0.20
0.23


1.07
0.07
0.98
1.17


The tetra-lamellate internal barrier is confined to the
penultimate and body whorls.
No close relationship is apparent between
Holospira fortisculpta and other known species. Most
of Puebla has been only superficially explored for mol-
lusks. The state is large with an area of 33,919 km2 and
has vast, but fragmented habitat suitable for Holospirinae.
It is not surprising that this and other disparate species-
groups of Holospira are known from so few localities.
Additional field work almost anywhere within the state
can be expected to yield additional populations and spe-
cies.
Etymology.- The name fortisculpta is from the
Latin fortis, meaning strong, and sculptus, a sculpture,
alluding to the strong sculpture characteristic of this spe-
cies.

The Holospira haploplax species-group
The group consists of a single species from cen-
tral Puebla. It is unique within Holospira by having a
single lamella comprising the internal barrier. It resembles
Bostrichocentrum (Figs. 165-168) in this trait, but the
resemblance is superficial. Bostrichocentrum have stout
white, opaque smooth or striate shells, and they lack a
columellar lamella as such (Thompson 1964). Instead,
they have a low hollow bulge around the base of the
columella in the last whorl (Fig. 168). At one time all
holospirids with a single lamella or lamella-like bulge were
placed in Bostrichocentrum. Thompson (1974) dem-
onstrated that Bostrichocentrum as previously under-
stood was polyphyletic and restricted the genus to in-
clude only those species from southern Mexico. On the
basis of the over-all shell morphology Holospira
haploplax strongly resembles species of Stalactella.
The latter group differs by having a strong columellar
lamella and a dentate parietal lamella.


Holospira haploplax n. sp.
Figs. 159-164
Description.- Shell opaque; light brown with
nearly white oblique ribs; peristome white; interior of
aperture light brown. Cylindrical-conical in shape; apex
relatively short and obtuse-convex; spire straight-sided.
Medium-sized and stocky; length about 10.9-13.9 mm;
0.24-0.31 times as wide as long. Whorls 13.4-15.6; apex
about 1/5 the length of the shell and consisting of about
seven whorls; apical whorls convex; whorls on spire flat-
sided with a distinct but shallow suture; embryonic whorls
2.0-2.3, smooth; subsequent whorls sculptured with
strong, oblique posteriorly arched ribs separated by
interspaces that are about 2-3 times as wide as ribs
Penultimate whorl with 32-54 ribs. Base of shell rimate
or imperforate. Aperture broadly auriculate; about as
wide as high; free from and projecting forward slightly
beyond previous whorl (Fig. 162); prosocline at an angle
of about 20 to shell axis. Peristome strong and moder-
ately reflected; narrowest at posterior corner; widest
along columellar margin. Columella about 0.1 times di-
ameter of shell, straight, hollow. Internal barrier mono-
lamellate, consisting of a single lamella (Figs. 163, 164);
parietal, basal and palatal lamellae absent; columellar
lamella strong; not visible from within aperture; originat-
ing at beginning of penultimate whorl and extending
through first half of body whorl; located just below cen-
ter of columella; projecting about a third of distance
across cavity.
Measurements of the holotype and 25 paratypes
(UF 337498) are given in Table 17.
Type Locality.- Puebla, 12.3 km east of Puebla,
2525 m alt. Holotype: UF 34381; collected 9 July, 1966
by Fred G. Thompson. Paratypes: UF 255392 (27),
ITCV (10); same data as the holotype. Other speci-
mens: UF 337498; same data as the holotype, preserved
in 75% ethanol.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Table 17. Holospira haploplax n. sp. Shell parameters of the holotype (UF 34381) and 25 paratypes (UF 337498).

L W AH AW Wh RP W/L AH/L AW/AH


Holotype
Paratypes
X
SD
Min.
Max.


12.7 3.4 2.4 2.3 15.0 32 0.27 0.19 0.97


12.23
0.84
10.9
13.9


3.45
0.10
3.4
3.6


2.44
0.23
2.3
2.9


2.41
0.18
2.3
2.9


14.35
0.78
13.4
15.6


39.58
5.65
32
54


0.28
0.02
0.24
0.31


0.20
0.02
0.17
0.24


0.99
0.09
0.82
1.13


The type locality is about 1 km south of Hwy 150
on a limestone hill, and was in an open oak grove with a
ground cover of grasses. Snails were found aestivating
on limestone boulders among grasses and leaves close
to the ground.
Distribution. Known only from the type locality.
Remarks.- The shell is brown with strong white
ribs, medium sized, stout, and cylindrical with a rela-
tively short obtuse apex. The ribs are oblique, stout and
about a third the width of their interspaces. The internal
barrier consists of a single strong lamella, the columellar
lamella, that is confined to penultimate whorl and the
first half of the body whorl.
Etymology. The species name haploplax is from
the Classical GreekpClrrog meaning single or simple,
andizAa4, something wide and flat. The name alludes
to the single lamella comprising the internal barrier.

Subgenus Stalactella Bartsch, 1906
Stalactella Bartsch, 1906: 151. Rehder, 1940:315. Pils-
bry, 1953: 135.
Type species.- Holospira (Stalactella) rose
Bartsch, 1906. Four other species are included in
Stalactella.
Stalactella includes a small group of species found
in Puebla and immediately adjacent northeastern
Oaxaca. The shell is opaque brown and bears low,
crowded ribs. The smooth embryonic whorls are turgid
and button-shaped and are about equal in size. The fol-
lowing half postembryonic whorl is wider but shorter,
offsetting the two teat-like earlier whorls at a slightly
oblique angle (Fig. 173a). The internal barrier is bi-lamel-
late, consisting of the columellar lamella and the parietal
lamella. The columellar lamella is smooth, thick and
wedge-shaped along its edge (Fig. 177). The parietal
lamella is thin, low, rounded and bears slender denticles
arranged in a single row. In some species the parietal
lamella is reduced to a thin callus. The penultimate whorl


is abbreviate and thick-walled (Figs. 176, 177, 181, 183,
190, 195). This is the section of the spire that bears the
principal portion of the internal barrier. Here the shell
wall is 3-4 times as thick as in the preceding or following
whorls. The abbreviate penultimate whorl with its thick
wall appear to be unique in Holospira.
Previous authors (Rehder 1940, Pilsbry 1953) treat
Stalactella as a section within Propilsbrya Bartsch,
1906 [Type species: Epirobia (Propilshba) nelsoni
Bartsch, 1906]. Both Propilsbrya and Stalactella have
fine pectinate denticles arranged in a single row along
the parietal lamella similar to Stalactella. This similar-
ity appears to be a matter of convergence. In
Stalactella the parietal and the columellar lamella are
confined to the lower most whorls of the spire, as is
characteristic of the basic arrangement in the Holospira,
and the parietal lamella bears a single row in elongate
denticles.
Propilsbrya is a disparate assemblage of species found
in Coahuila, Tamaulipas and San Luis Potosi. It includes
three subgenera. Propilsbrva (s.s.) is bi-lamellate. The
internal barrier consists of a columellar lamella and a
parietal lamella. The columellar lamella is restricted to
the whorls of the spire and is smooth. The parietal lamella
extends through nearly the length of the apex and the
spire, and its edge is serrate. Pectinistemma Rehder,
1940 also is bi-lamellate. The lamella are confined to
the lower whorls of the spire, and both the columellar
lamella and the parietal lamella are serrate.
Prionoloplax Pilsbry, 1953 is tri-lamellate with the bar-
rier confined to the penultimate and anti-penultimate
whorls. The columellar lamella and the basal lamella
are smooth. The parietal lamella is serrate.
Propilsbrya potosiana Pilsbry, 1953 from Nuevo
Le6n was originally placed in Stalactella because it has
a columellar lamella and a dentate parietal lamella.
Pilsbry (1953) treated Stalactella as a section within
Propilsbrya. The similarities in lamellar structure be-







tween P potosiana and Stalactella appears to be due
to convergence, not phylogeny. The species more
strongly resembles typical species of Propilsbrya than
Stalactella by having an elongate-tapered apex, the
embryonic whorls are not offset as they are in
Stalactella, and the penultimate whorl is not abbrevi-
ated and thick-walled.
The development of serrate or dentate lamella
within mainland urocoptids occurs in several evolution-
ary lines, including Propilsbrya, Pectinostemma,
Holospira acanthidia, Coelocentrum, subgenus
Ptychodonta Bartsch, 1906, and Anisospira, subgenus
Trachycion Thompson, 1968 (Thompson, 1968; Thomp-
son & Correa-Sandoval, 1994). Until the systematics
of the subgenera of Propilsbrya are better understood
their relationships to Stalactella remains unresolved.

Holospira (Stalactella) rosei Bartsch, 1906
Figs. 169-171
Holospira (Stalactella) rosei Bartsch, 1906: 151-153;
pl. 4, fig. 10.
Description.- The holotype (USNM 188181) and
three fragmentary paratypes (USNM 188181 a) are the
only known specimens, all of which are broken. The
following description is based on this material. The faded
shells suggest that the live shell is light brown. Medium
sized, broken holotype 14.4 mm long, 3.7 mm. wide, and
less than 0.26 times as wide as high. Shell elongate-
cylindrical; apex attenuated, about half length of the shell.
Suture around apex moderately impressed so that the
whorls appear to be slightly telescoped. Last two whorls
of the spire slightly narrower than previous whorls.
Penultimate whorl shorter than preceding whorl. Last
whorl enlarged and indented with a shallow longitudinal
furrow below periphery so that neck is triangular in cross
section with a narrowly rounded base. Neck of last
whorl projecting forward for about 1/4 of diameter of
shell (Fig. 170). Umbilicus open but narrowly perfo-
rate. Sixteen whorls remaining in holotype; about 3-4
earliest whorls are missing due to unnatural break, sug-
gesting a species with about 19-20 whorls; remaining
apex consisting of 8 whorls which are strongly arched
between the suture. Whorls of spire flattened, with a
moderately impressed suture. Apex has numerous, ob-
lique, fine ribs that are as high as wide and are about
half as wide as their interspaces. Ribs uniformly wide
throughout their lengths. Ribs become vague and poorly
defined on middle of spire, but becomes stronger and
more clearly defined on last two whorls. Ribs continu-
ous around neck of last whorl. Aperture trapezoidal in
outline, 2.3 mm wide and 2.4 mm high, broadly auricu-
late in shape internally because of an external indenta-


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

tion into parietal wall; aperture 0.96 times as wide as
high. Width of aperture 0.62 times width of shell. Pari-
etal lip nearly straight. Peristome relatively thin and
weakly reflected along outer, basal, and lower columel-
lar lip; less strongly reflected along parietal and upper
columellar lip. Plane of aperture in lateral profile paral-
lel to axis of shell (Fig. 170). Axis hollow, about 0.1
times width of shell. Internal barrier consists of two
lamellae (Fig. 171). Columellar lamella extending from
anti-penultimate whorl to base of last whorl; its lower
end visible within aperture. Columellar lamella thick and
rounded along its outer edge; transversely sloping slightly
downward. Parietal lamella 2.3 whorls long; originating
in penultimate whorl and extending downward to last
quarter of body whorl. Parietal lamella very weakly
developed in holotype as a thin narrow ridge on middle
of septum. Parietal lamella bearing a series of sharp
denticles that are about as long as their interspaces and
are curved outward. Approximately 30 denticles are
present in penultimate whorl.
Type Locality.- Puebla, "Tehuacan". Holotype:
USNM 188181; collected by Joseph N. Rose and Jo-
seph H. Painter. Paratypes: USNM 188181 a (fragments
of 3 specimens).
Distribution.- Unknown. This species is reported
to have been collected at Tehuacan, but the exact prov-
ince is uncertain. We have found no populations in the
vicinity of Tehuacin that resemble this species.
Remarks.- This is a species of Stalactella that is
recognized by its dull brown shell with an attenuated
apex. The apex and base of the shell are sculptured
with numerous fine retractive riblets that are absent from
the middle of the spire.
The dull brown color and the sculpture are most
like the following species. The two differ in that with
Holospira rosei the apical whorls are telescoped, as
opposed to being scalariform. The attenuated apex of
H. rosei is unlike the short abbreviate apex characteris-
tic of other Stalactella. It is distinguished also from
other known species by it numerous fine closely spaced
ribs on the apex and on the lower part of the spire. The
penultimate whorl has about 49 ribs whereas other spe-
cies have fewer ribs.
Bartsch (1906:153) described this species as be-
ing decollated. Rehder(1940:315) and Pilsbry (1953:135)
repeated this information. The apical whorls in the ho-
lotype are not decollated. They were broken from the
shell subsequent to death, for there is no internal apical
plug as would be formed if the shell was alive when the
whorls were lost. The shells of all other Stalactella that
we have examined have complete apexes as is typical
of the Holospirinae. We know of no examples in the





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

subfamily Holospirinae in which the apex is not normally
complete.

Holospira (Stalactella) cremnobates n. sp.
Figs. 172-177
Description.- Periostracum uniform light brown
with lighter colored ribs. Ribs on last one or two whorls
usually white. Peristome white; interior of aperture light
brown. Medium sized for subgenus, 11.6-14.1 mm long;
0.25-0.31 times as wide as long. Shell slightly club-shaped
or nearly cylindrical. Widest at base of apex, and nar-
rows toward last whorl, which is noticeably enlarged.
Apex short and convex; with 3.0 button-shaped, strongly
protruding mammaform embryonic whorls (Fig. 173a).
Apical whorls weakly scalariform, not telescoped. Last
whorl noticeably enlarged and with a short neck that in
lateral profile projects forward beyond last whorl for
about 1/6 diameter of shell (Fig. 173). Neck with a
broad, weakly impressed longitudinal furrow below pe-
riphery. Dorsum of neck with an impressed longitudinal
furrow that creates an elongate parietal denticle inter-
nally. Umbilicus open as a small, nearly round hole.
Penultimate whorl shorter than preceding whorl and fol-
lowing whorls (Figs. 172, 176). Suture strongly impressed
on apex and weakly impressed on spire. Shell with 15.0-
17.7 whorls. Lower whorls of apex weakly arched.
Upper whorls of spire flattened. Apex consists of about
9 whorls that are separated by a deeply impressed su-
ture and are strongly rounded at the periphery. Embry-
onic shell consisting of about 3 smooth, strongly protrud-
ing whorls. First two embryonic whorls enlarged and
button-shaped. The third embryonic whorl is noticeably
shorter but wider. Following apical whorls have strong
oblique ribs that are very narrow at their upper ends and
increase in height and width toward their bases. The
ribs are about 1/3 the width of their interspaces. Spire
with weaker but distinct reversed-sigmoid ribs that are
1/3-1/5 the width of their interspaces. Ribs becoming
stronger on penultimate and last whorls. They continue


onto neck where they diminish in width on base and
inner edge of neck. Penultimate whorl with 27-42 ribs.
Aperture broadly auriculate in shape, and projecting for-
ward beyond preceding whorl (Fig. 173). Aperture width
about equal to or slightly greater than aperture height,
and about 0.7 times the width of the shell. Peristome
moderately thickened and nearly uniformly reflected
around aperture. Plane of aperture slightly oblique to
axis of shell. Axis straight, hollow, and slightly concave
below middle of whorls. Axis about 0.14-0.17 times
width of shell. Internal barrier consists of two lamellae,
the columella lamella and the parietal lamella, which are
of about equal length and locations within the shell (Fig.
177). Columellar lamella not visible within aperture.
Beginning in anti-penultimate whorls and extends down-
ward to halfa whorl behind aperture. Columellar lamella
thick and wide in last two whorls; its outer edge forming
a thickened cord along edge of lamella; extending about
1/3 of distance across width of cavity; without a notice-
able outward slope. It then rapidly decreases in size to
its upper termination. Parietal lamella a low narrow ridge
located in anti-penultimate and penultimate whorls about
2/3 of distance from columella to outer wall; about 1.5-
2.0 whorls long, extending downward to half a whorl
behind aperture. It bears a row of fine denticles that
curve outward. Denticles variable in size but are equal
in length to or are slightly shorter than their interspaces.
Five specimens have 21,23, 25, 26 and 27 denticles re-
spectively.
Measurements of the holotype and 25 paratypes
(UF 21453) are given in Table 18.
Type Locality.- Puebla, 10 km north of Tehuacan
(180 37' 23" N, 970 23' 35" W); 1990 m alt. Holotype:
UF 190789; collected 2 April, 1991 by Fred G. Thomp-
son and Elizabeth L. Mihalcik. Paratypes: UF 21453
(91), same data as the holotype; UF 34300 (107), ITCV
(60), same locality, collected 8 August, 1965 by F. G.
Thompson.
The type locality is on the north side of a canyon


Table 18. Holospira (Stalactella) cremnobates n. sp. Shell parameters in mm for the holotype (UF 190789) and
25 paratypes (UF 21453).


W AH AW


Wh RP W/L AH/L AW/AH


13.0 3.6 2.4 2.3 16.5 27 0.28 0.18 0.94


12.9
0.72
11.6
14.1


3.6
0.16
3.3
4.1


2.5
0.15
2.3
2.8


16.2
0.78
15.0
17.7


32.8
4.83
27
42


0.28
0.19
0.25
0.31


0.18
0.02
0.15
0.22


1.04
0.07
0.94
1.20


Holotype
Paratypes
X
SD
Min.
Max.







about 0.5 km east of Hwy 150. The area consists of a
series of bare limestone cliffs with long talus slopes. A
dense forest of submesic scrub covered the valley floor
below the cliffs. The tops of the cliffs were covered
with grasses. Holospira cremnobates was collected
only on the face of nearly vertical limestone cliffs from
under spalled slabs of limestone and clusters of dried
mosses. Bostrichocentrum sp. was abundant in the
tallus slopes below the cliffs.
Distribution.- Known only from the type locality.
Remarks.- Holospira cremnobates is most simi-
lar to the following species, H. (S.) psectra. It is distin-
guished from the latter by its club-shaped shell sculp-
tured with thinner, lower, sparser ribs that expand only
slightly at their bases. The columellar lamella is longer
and much stronger, bearing a cord-like thickening along
its outer edge. The parietal lamella is a ridge that lies
about 1/3 of the distance from the outer wall, and bears
denticles that are about equal to or slightly shorter than
their interspaces.
Etymology.- The species name cremnobates is
from the Greek Kpe voparfq, meaning a frequenter
of steep slopes, and alludes to the habitat of this species.

Holospira (Stalactella) psectra n. sp.
Figs. 178-183
Description.- Periostracum uniform light brown
in color, usually with ribs. Peristome white: interior of
aperture light brown. Shell medium-sized for subgenus,
11.5-14.0 mm long; and 0.26-0.31 times as wide as high.
Slightly club-shaped, widest at base of apex, and nar-
rows toward last whorl. Apex relatively short, and coni-
cal. Last whorl noticeably enlarged, and has a short
neck so that in lateral profile aperture hardly projects
forward beyond the periphery of the preceding whorl
(Fig. 179). Last quarter whorl constricted behind aper-
ture and pinched inward below periphery. Umbilicus
open but narrow. Penultimate whorl slightly shorter than
preceding whorl. Whorls 15.1-18.1. Apex consisting of


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

about 9 strongly rounded, slightly scalariform whorls that
are separated by a deeply impressed suture. Embry-
onic shell consisting of about 2-3 smooth, strongly pro-
truding, button-like whorls. Last embryonic whorl no-
ticeably enlarged. Following whorls of spire weakly
arched; Apex sculptured with very strong oblique ribs
that increase in height and width towards their bases,
and are about 1/3 width of their interspaces. Spire sculp-
tured with slightly weaker but distinct rib that are 1/3-1/
5 width of their interspaces and are almost knob-like at
their bases. Ribs become stronger on penultimate and
last whorls and continue onto neck where they diminish-
ing in width. Penultimate whorl with 35-47 ribs. Aper-
ture broadly auriculate in shape. Aperture width about
equal to or slightly greater than its height, and about 0.7
times width of shell. Peristome moderately thickened
and nearly uniformly reflected. Plane of aperture slightly
oblique to longitudinal axis of shell. Axis hollow and
about 0.14-0.17 times width of whorl. Internal barrier
bi-lamellate. Columella lamella (Fig. 183) begins in lower
half of anti-penultimate whorl and extends downward to
0.5 whorls behind aperture; not visible within aperture.
It lies about a third of distance above the floor of whorl.
It is thickest and widest in the penultimate whorl and
projecting outward for less than 1/4 of width of cavity
without a noticeable downward slope; it is low and
weakly developed in the last whorl. Parietal lamella a
narrow weak spiral callus on parietal wall about a third
of distance from outer wall. It begins in the lower half
of anti-penultimate whorl and extends downward to
about a half whorl behind aperture, although usually
shorter then columellar lamella. It bears a row of fine
short denticles that curve outward. Denticles variable
in size, but usually less than half the length of their
interspaces. Five specimens have 14, 18, 20, 21 and 22
denticles respectively.
Measurements of the holotype and 24 paratypes (UF
233197) are given in Table 19.
Type Locality.- Puebla, a limestone cliff on the


Table 19. Holospira (Stalactella) psectra n. sp. Shell parameters in mm for the holotype (UF 233197) and 25
paratypes (UF 41472).


L W AH AW Wh RP W/L AH/L AW/AH

12.4 3.7 2.2 2.2 16.0 36 0.30 0.28 0.97


12.6 3.7
0.6 0.1
11.5 3.5
14.0 3.9


2.30
0.1
2.2
2.6


2.30
0.1
2.2
2.4


16.5
0.6
15.1
18.1


0.29
0.01
0.26
0.31


0.18
0.01
0.16
0.20


1.01
0.30
0.97
1.10


Holotype
Paratypes
X
SD
Min.
Max.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

south slope of Cerro Caolalote, 2 km east of El Carmen
(18 36"N, 9726'01"W); 1870 m altitude. The type
locality is at the base of a south-facing bare limestone
cliff. Live specimens were found under limestone rubble
and dead Agave. Holotype: UF 233197; collected 25
October, 1992 by Fred G. Thompson and Elizabeth
Mihalcik. Paratypes: UF 41472 (106), ITCV (50); same
data as the holotype.
Distribution.- Known only from the type locality.
Remarks.- This species is most closely allied to
but differs from Holospira cremnobates by its nearly
cylindrical-shaped shell in which the penultimate whorl
is only slightly shorter than the preceding whorl. The
ribs are white, more numerous, and stronger, and are
almost knob-like at their bases. The columella lamella is
weaker, and slopes outward slightly from the columella.
The parietal lamella is reduced to a thin spiral callus and
bears short, conical denticles that are less than half the
length of their interspaces.
Etymology.- The species name psectra is from
the Classical GreekspKT-pameaning a currycomb, and
alludes to the comb-like parietal lamella.

Holospira (Stalactella) marmorata n. sp.
Figs 184-190
Description.- Shell white with numerous irregu-
lar hydrophanus zones that form a marbled appearance.
Ribs predominantly white in contrast to darker intern-
odes and hydrophanus blotches. Embryonic whorls and
early whorls of apex dark tan with lighter colored ribs.
Peristome white; interior of aperture light rust-colored.
Shell medium-sized, about 12-15 mm long; 0.24-0.29
times as wide as high; nearly cylindrical in shape; wid-
est just above middle of shell and tapering slightly be-
low. Apex is moderately short and convex with protrud-
ing embryonic whorls. Penultimate whorl shorter than
preceding whorl. Last whorl greatly enlarged, and usu-
ally has a slightly longitudinal constriction below periph-
ery. Last whorl with a short neck that projects forward
about 1/5-1/6 diameter of shell (Fig. 185). Base of last
whorl rounded. Umbilicus rimate, with a very narrow
perforation. Suture moderately impressed with weakly
arched whorls. Whorls 15.5-18.9. Apex consists of
about 9-10 moderately arched whorls. Embryonic shell
consisting of 2.5 smooth, protruding, button-shaped
whorls. Following apical whorl considerably shorter but
wider than embryonic whorls. Subsequent whorls of
spire weakly arched, but surface curvature accentuated
by ribbed sculpture. Sculpture of apex consisting of
moderately strong ribs that are about 1/2-1/3 width of
interspaces. They become wider, node-like and crenu-
late lower suture. Ribs on the middle of spire lower but


clearly defined and are about 1/3-1/4 width of
interspaces. Ribs become stronger and higher on last
two whorls, and then become weaker towards peris-
tome. Penultimate whorl with 29-55 ribs. Aperture
broadly auriculate in shape, with a strong longitudinal
indentation on parietal wall. Aperture about 0.88-1.12
times as high as wide, and about 0.15-0.20 times the
length of shell. Peristome thickened and nearly uniformly
reflected around aperture. Plane of the aperture in lat-
eral profile parallel or slightly oblique to the shell axis
(Fig. 185). Internal barrier bi-lamellate (Fig. 190); not
visible from within aperture. Columellar lamella two
whorls long; beginning abruptly in penultimate whorl and
ending at 2 whorl behind aperture; located just below
middle of columella and sloping outward about half-way
across cavity; conspicuously thickened along its middle
and narrows to its outer edge forming a channel along
columella on its upper side and a deep concavity below.
Parietal lamella a low callus located along outer third of
the roof, and is about halfa whorl longer than columellar
lamella. It begins in anti-penultimate whorl and ends a
half whorl behind aperture. Parietal lamella with 17-22
denticles arranged in a single row along its crest. Den-
ticles are about as long as their interspaces and curve
outward.
Measurements of the holotype and 25 paratypes
(UF 80954) are given in Table 20.
Type Locality.- Puebla, 13.5 km WSW of San
Bartolo Teontepec (18" 26' 19" N, 970 36' 42" W); 2120
m alt. The area is a xeric limestone hillside on the north
side of the highway. Snails were found under caliche
slabs and limestone boulders. Holotype: UF 233188;
collected 21 February, 1992 by Fred G. Thompson.
Paratypes: UF 80954 (30); same data as holotype.
Distribution.- Puebla: 6 km N of Santa Cruz
Nueva (18 20' 19" N, 970 49' 35" W), 1760 m alt. (UF
200956); 4.5 km S of San Vicente Coyotepec (180 20'
19" N, 970 49' 59" W) 2080 m alt. (UF 200601); 3.7 km
S of San Vicente Coyotepec, 1880 m alt. (UF 233190,
ITCV).
Remarks. Distinguished by its low dome-shaped
apex. The enamel-like white shell is marked with ir-
regular gray hydrophanus blotches producing a marbled
color pattern. Columellar lamella is thickened in its cen-
ter to form a channel above along columella and below
a concavity. The parietal lamella bears 17-22 denticles
that are about as long as their interspaces and curve
outward.
This species shows considerable local variation in
size. Specimens from near Coyotepec are smaller than
typical, but there is broad overlap in all measurements
between the various populations. A series of specimens





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Table 20. Holospira (Stalactella) marmorata n. sp. Shell parameters in mm for the holotype (UF 233188) and 25
paratypes (UF 80945).


L W AH


AW Wh RP W/L AH/L AW/AH


13.9 3.8 2.6 2.5 16.0 32 0.27 0.19 0.96


13.4
0.87
11.9
15.2


3.6
0.12
3.4
3.8


2.4
0.16
2.2
2.7


2.4
0.19
2.1
2.7


16.9
0.85
15.5
18.9


34.9
6.3
28
53


0.25
0.03
0.24
0.29


0.18
0.02
0.15
0.20


1.02
0.09
0.88
1.12


from 3.7 km south of Coyotepec vary in length from
10.1-13.6 mm and have 13.6-17.4 whorls with 35-54
ribs on the penultimate whorl. Other aspects of the shell,
both external and internal, are typical.
Etymology.- The species name is from Latin,
marmorata, meaning marbled, and alludes to the color
of the shell.

Holospira (Stalactlla) chazumbae n. sp.
Figs.191-195
Description.- None of the specimens compris-
ing the type lot were live collected. However, many are
fresh shells. Ground color enamel white with extensive
hydrophanus zones on apex and spire. Embryonic whorls
and upper half of apex tan colored, but with white ribs
appearing on the postembryonic whorls. Peristome
white; interior of the aperture rust-colored. Shell 12.5-
17.3 mm long, and 0.21-0.30 times as wide as long. It is
cylindrical or slightly club-shaped, widest just above
middle. Apex relatively attenuated for subgenus. Em-
bryonic whorls about 2.5, protruding, button-like; offset
from the spire by first halfpostembryonic which is shorter,
but wider. Penultimate whorl shorter than preceding
whorl. Body whorl conspicuously enlarged with a short
neck that is about 1/6 diameter of preceding whorl (Fig.
192, 193). Neck with a weak longitudinal furrow below
periphery. Base narrowly rounded. Umbilicus rimate.
Suture on spire moderately impressed between weakly
arched whorls. Whorls 16.3-20.4. Apex attenuated,
consists of 11-12 weakly arched whorls. Apical sculp-
ture with moderately oblique ribs that are about half as
wide as their interspaces. On middle of spire ribs tend
to become obsolete, and then become strong and well
defined on last two whorls. Penultimate whorl with about
32-50 ribs. Aperture broadly auriculate in shape with a
strong longitudinal indentation protruding into parietal
margin; aperture about 0.96-1.20 times as high as wide,
and is about 0.12-0.19 times length of shell. Posterior


comer of aperture noticeably narrower and more sharply
angular than in related species. Peristome thickened
and moderately expanded; narrowest along posterior
corer and widest along baso-columellar margin. Plane
of aperture in most specimens nearly parallel to shell
axis in lateral profile; occasionally sloped downward in
some specimens. Axis is straight and hollow, and is about
0.15-0.19 times the width of the whorl. Internal barrier
bi-lamellate (Fig. 195). Columellar and parietal lamel-
lae, both extend to about 0.6 whorls behind the aperture;
not visible from within aperture. Columellar lamella 2.5
whorls long and begins abruptly in anti-penultimate whorl;
thickened along its middle, and slightly thinner along its
periphery; widest and strongest in penultimate whorl
where it projects slightly downward about half way
across cavity and forms a concavity on its ventral sur-
face along the columella. Parietal lamella a low callus
on center of dorsal wall; with sparse, widely spaced
denticles that curve outward and slightly forward. Three
specimens have 19, 16 and 16 teeth respectively in
penultimate whorl.
Measurements of the holotype and 24 paratypes
(UF 214439) are given in Table 21.
Type Locality.- Oaxaca, hillside 2 km northeast
of Santiago Chazumba; 1970 m alt. Snails were col-
lected from under blocks of caliche among sparse
growths of xeric scrub vegetation. Holotype: UF 34299;
collected 20 October, 1970 by Fred G. Thompson.
Paratypes: UF 214439 (80), ITCV (30); same data as
the holotype. This species was found in association with
Bostrichocentrum sp.
Distribution.- Known only from the type locality.
Remarks.- The shell is similar to Holospira
marmorata because of its enamel-like white color
marbled with hydophanus blotches and streaks. It dif-
fers from H. marmorata in that the apex is more at-
tenuate, approaching that of H. rose. Holospira
marmorata has a short dome-shaped apex. In H.


Holotype
Paratypes
X
SD
Min.
Max.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Table 21. Holospira (Stalactella) chazumbae n. sp. Shell parameters in mm for the holotype (UF 34299) and 24
paratypes (UF 214439).

L W AH AW Wh RP W/L AH/L AW/AH


Holotype
Paratypes
X
SD
Min.
Max.


14.1 3.6 2.3 2.2 18.2 30 0.25 0.16 0.96


3.7
0.23
3.5
4.0


2.4
0.29
2.2
3.7


18.1
0.99
16.6
20.5


36.7
4.1
30
46


0.26
0.02
0.21
0.30


0.17
0.12
0.15
0.19


0.99
0.12
0.81
1.40


chazumbae the posterior corer of the aperture is no-
ticeably narrower and more sharply angulate than in re-
lated species.
Etymology.- The species name chazumbae is a
patronymic taken from the Village of Santiago Chazumba,
near the type locality.

Genus Coelostemma Dall, 1895
The following species is included in this account in
order to clarify the identity and status of a name that has
been in question since it was proposed.

Coelostemma microstoma (Pfeiffer, 1861)
Figs. 196-198
Cylindrella microstoma Pfeiffer, 1861: 27. Pfeiffer,
1862a: 81. Pfeiffer, 1868: 390.
Holospira microstoma (Pfeiffer). Fischer & Crosse;
1873: 337; pl. 17, figs. 9-9a. Martens, 1897: 287.
Pilsbry, 1902: 102-103; pl. 15, figs. 4-5.
Coelostemma (?) microstoma (Pfeiffer).- Bartsch, 1943:
58.
Description.- Color uniform opaque white; lec-
totype lusterless, suggesting that the specimen had been
dead and sun-bleached before it was collected. Shell
medium sized for the genus, 15.3 mm long, and 1/3 as
wide as high; club-shaped and with aperture projected
forward on a short neck that is about 1/4 the diameter of
penultimate whorl (Fig. 197). Umbilical perforation round
and about 1/8 the width of last whorl. Lectotype with
17.8 whorls. Whorls of spire nearly flat-sided and sepa-
rated by a weakly impressed suture. Apex dome shaped
with weakly arched whorls that tend to be slightly sca-
lariform. Embryonic shell containing 2.0 smooth whorls;
first whorl protruding and button-shaped; second whorl
very short and wider. Following apical whorl distinctly
ribbed, which over the next four whorls fade into in-
creasingly finer protracted vertical striations. Spire
smooth, except for the last whorl, which has weak low


rounded ribs. Ribs begin below the periphery and be-
come increasingly stronger, more crowded and higher
on last half of body whorl and neck. Axis large and
hollow, 0.35 times width of spire; concave within each
whorl (Fig. 198). Aperture about as high as wide; au-
riculate in shape with a strong impression on dorsal wall
behind the peristome. Interior of aperture sub-ovate in
shape; columellar wall nearly flat and vertical. Dorsal
and outer lip thickened and weakly reflected; basal lip
reflected, but not conspicuously thickened; columellar
lip broken in lectotype.
Measurements of the lectotype are as follow:
length, 15.3 mm, width, 5.1 mm; aperture height, 2.5
mm; aperture width, 2.5 mm.
Type Locality.- Unknown. Lectotype by present
designation: British Museum (Natural History) 1996162.
The specimen was labeled "Syntype ?". It matches
very closely the description, measurements and whorl
count given by Pfeiffer (1861:27) to such a degree that
there is little doubt that it is the specimen upon which the
original description was based. Also, it is consistent with
the figures made by Sowerby and published in Fischer
& Crosse (1873: pl. 17, figs. 9-9a.), except that their
figures show distinct riblets on the last whorl. It is not
clear from Pfeiffer's description how many specimens
he had examined. I have been unable to locate others.
Distribution.- Unknown.
Remarks.- A species distinguished by its me-
dium size club-shaped shell, its dome-shaped apex that
is nearly uniformly rounded, and its sculpture, which is
ribbed on the earliest postembryonic whorls and then
becomes increasingly finer over the surface of the apex;
the whorls of the spire are smooth, except for the ribbed
last whorl.
Coelostemma microstoma appears to be most
closely related to C. herrerae (Bartsch, 1906) from
Silaca Yoapan (= Silacayoapan), Oaxaca, because of
their uniformly rounded dome-shaped apexes, their similar







sculpture and their claviform shapes. Coelostemma
herrerae is a stockier species, being 0.35-0.44 times as
high as wide, the columella is almost half the diameter
of the shell, the columella is nearly straight sided, not
concave as in C. microstoma, and it bears a low spiral
swelling below the middle in the last three whorls.

ACKNOWLEDGMENTS
The following people have loaned specimens to us from
the collection in their charge. Dr. Matthias Glaubrecht,
Museum fur Naturkunde, Hamberg Universtat zu Ber-
lin, Germany. Dr. Ronald Janssen, Senkenbergische
Naturforschenden Gesellschaft, Frankfurt, Germany.
Fred Naggs, British Museum (Natural History) London,
UK. Dr. Robert Hershler, National Museum of Natural
History, Washington, DC. We thank Anthony B. Falcetti
and Heather Walsh-Haney of the C. A. Pound Labora-
tory, University of Florida, Gainesville, Florida for radio-
graphs of the type specimens of Cylindrella goniostoma,
Cylindrella teres, Holospira hogeana, Cylindrella
pfeifferi, and Cylindrella microstoma. We thank John
Slapcinsky, Florida Museum of Natural History, for as-
sistance with the many tasks that were essential to the
preparation of this report. Kenneth Emberton and Ned
Strenth provided helpful reviews of this paper. Gregg
Brewer, Fredericksburg, Texas. provided valuable field
assistance in Mexico. Especially, we thank Alfonso
Correa-Sandoval, Instituto Tecnologico de Cd. Victoria,
Tamaulipas for his assistance with field work and for
many other courtesies that have enhanced this project.
Finally, we are grateful for reviews by Ned E. Strenth,
Kenneth Emberton, and an anonymous reviewer. Their
attention to detail have made this a better paper. Col-
lecting permits for Mexico were issued by the Direccion
General de Fauna Silvestre (1966-1991) and by the
Direccion General de Vida Silvestre. 1995-2002.

LITERATURE CITED
Albers, J. C. 1860. Die Heliceen nach nattirlicher
Verwandtschaft systematisch geordnet. 2:1-359.
Edition edited by E. von Martens. Leipzig.
Bartsch, P. 1906. The urocoptid mollusks from the
mainland of America in the collection of the
United States National Museum. Roceedings
of the United States National Museum, 31: 109-
160; pls. 3-5.
Bartsch, P. 1926. New urocoptid land shells from
Mexico. Proceedings of the United States Na-
tional Museum, 70: 1-13; pl. 1.
Bartsch, P. 1945. New urocoptid mollusks from Mexico.
Journal of the Washington Academy of Sciences,
35: 92-95.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)

Bartsch, P. 1947a. Some Mexican Urocoptid mollusks.
Journal of the Washington Academy of Sciences,
37:141-142.
Bartsch, P. 1947b. Notes on some Mexican urocoptid
mollusks, with the description of new species.
Journal of the Washington Academy of Sciences,
37: 284-288.
Bequaert, J. E., & W. B. Miller. 1973. The mollusks of
the arid southwest with an Arizona check list.
University of Arizona Press. 1-271. Tucson.
Cockerell, T. D. A. 1914. Tertiary Mollusca from New
Mexico and Wyoming. Bulletin of the Ameri-
can Museum Natural History, 33: 101-107.
Fischer, P., & H. Crosse. 1870-1878. Mission
Scientifique au Mexique et dans L'Amerique
Central. Etudes sur les mollusques terrestres
et fluviatiles du Mexique et du Guatemala, 1: 1-
702; pl. 1-31. Paris.
Gilbertson, L. H. 1989a. A new species of Holospira
(Gastropoda: Pulmonata) from Sonora, with the
reproductive anatomy of Holospira minima.
Veliger, 32: 91-94; figs. 1-3.
Gilbertson, L, H. 1989b. A new species of Holospira
from Arizona, with the reproductive anatomies
of H. arizonensis and H. chiricahuana. Ve-
liger, 32:308-312.
Gilbertson, L. H. 1993. Reproductive anatomies of
Holospira spp. (Gastropoda: Pulmonata:
Urocoptidae) from Arizona and Sonora with a
new subgenus and a new subspecies. Ameri-
can Malacological Bulletin, 10: 71-81; figs. 1-
12.
Gilbertson, L. H., & E. Naranjo-Garcia. 1998. A new
subgenus and a new species of Holospira (Gas-
tropoda: Pulmonata: Urocoptidae) from Sonora,
Mexico. Veliger, 41: 314-318.
Gilbertson, L. H., & E. Naranjo-Garcia. 2004.
Millerspira, a replacement name for Millerella
Gilbert and Naranjo-Garcia, 1998. Veliger, 47:
157
Haas, F. 1933. Kurze Bermerkungen. Archiv ftir
Molluskenkunde, 65: 271.
Martens, E., in Albers, J. C. 1860. Die Heliceen, nach
nattirlicher Vervandtschaft systematisch
geordnet. Edit. 2. Edited by E. von Martens: 1-
359. Berlin.
Martens, E. von. 1890-1901. Biologia Centrali-Ameri-
cana. Mollusca. 1-706. British Museum (Natu-
ral History).
Menke, K. T. 1847. Vier neue Arten der Gattung
Cylindrella Pr. Zeitschrift fir Malakozoologie:





THOMPSON and MAHALCIK: Southern Mexican holospirid snails

Opinion 1932: International Commission of Zoological
Nomenclature. 1999. Holospira Martens, 1860
(Mollusca, Gastropoda): Cylindrella goldfussi
Menke, 1847 designated as the type species.
Bulletin of Zoological Nomenclature, 56: 206-
207.
Pfeiffer, L. 1841. Symbolae ad Historium Heliceorum:
1-88. Casselis.
Pfeiffer, L. 1848. Monographia heliceorum viventium,
2: 1-594. Liepsig. F. A. Brockhaus.
Pfeiffer, L. 1859. Monographia heliceorum viventium,
4: 1-920. Liepsig. F. A. Brockhaus.
Pfeiffer, L. 1861. Descriptions of forty-seven new spe-
cies of land-shells from the collection of H.
Cuming, Esq. Proceedings of the Zoological
Society, 1861: 20-29.
Pfeiffer, L. 1862a. Diagnosen neuer Heliceen.
Malakozoologische Blatter 8 [1861]: 77-84.
Pfeiffer, L. 1862b. Systematisches Conchylien-Cabi-
net: Die Gattung Cylindrella. 1-80; pls. 1-9.
Pilsbry, H. A. 1902-1903. Manual of Conchology, Ser.
II, 15:i-viii, 1-323; pls. 1-65.
Pilsbry, H. A. 1946. Land Mollusca of North America.
Monographs of the Academy of Natural Sci-
ences of Philadelphia. 3,2 (1): 1-520.
Pilsbry, H. A. 1953. Inland Mollusca of northern Mexico.
II. Urocoptidae, Pupillidae, Strobilopsidae,
Valloniidae, and Cionellidae. Proceedings of the
Academy of Natural Sciences of Philadelphia,
105: 133-167; pls. 3-10; text figs. 1-3.
Philippi, R. A. 1842-1847. Abbildungen und
beschreibungen neuer oder wennig bekannter
Conchylien, (I): 1-204. Cassel.
Rehder, H. A. 1940. A new urocoptid mollusk from
Mexico. Journal of the Washington Academy
of Science, 30: 315-316.
Strebel, H., & G. Pfeffer. 1880. Beitrag zur Kenntniss
der Fauna mexikanischer Land- und Siisswasser-
Conchylien. Theil IV: 1-122; pls. 1-15. Ham-


burg, J. J. Kerbst.
Thompson, F. G. 1964. Systematic studies on Mexican
land snails of the genus Holospira, subgenus
Bostrichocentrum (STYLOMMATOPHORA,
UROCOPTIDAE). Malacologia, 2: 131-143.
Thompson, F. G. 1968. Some Mexican land snails of
the family Urocoptidae. Bulletin of the Florida
State Museum, 12:125-183; fig. 1-29.
Thompson, F. G. 1971. Some Mexican land snails of
the genera Coelostemma and Metastoma
(Urocoptidae). Bulletin of the Florida State Mu-
seum, 15: 267-302; figs. 1-12.
Thompson, F. G. 1974. A xeric landsnail from New
Mexico. Southwestern Naturalist, 19: 53-56.
Thompson, F. G. 1988. The hollow-ribbed land snails of
the genus Coelostemma of the southwestern
United States and Mexico. Bulletin of the Florida
Museum of Natural History, 33: 87-111; figs. 1-
53
Thompson, F. G. 1998. Holospira Martens, 1850 (MOL-
LUSCA, GASTROPODA, UROCOPTIDAE):
proposed designation of Cylindrella goldfussi
Menke, 1947 as the type species. Bulletin Zoo-
logical Nomenclature, 55: 87-87.
Thompson, F. G, & A. Correa-[Sandoval]. 1991.
Mexican land snails of the genus
Hendersoniella. Bulletin of the Florida Mu-
seum of Natural History, 36: 1-23.
Thompson, F. G., & A. Correa-[Sandoval]. 1994. Land
snails of the genus Coelocentrum from north-
eastern M6xico. Bulletin of the Florida Mu-
seum ofNatural History, 36: 141-173.
Tryon, G. 1868. Monograph of the terrestrial Mollusca
of the United States. American Journal of Con-
chology, 3: 306-327.
Tozer, E. T. 1956. Uppermost Cretaceous and Pale-
ocene non-marine molluscan faunas of western
Alberta. Geological Survey of Canada, Mem-
oir 280: 1-125.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 2-10. Holospira. Figures 2-7. Holospira goniostoma (Pfeiffer, 1856). Lectotype (British Museum,
Natural History 1996153). Figures 2-5: Radiographs showing the positions of the internal lamellae. Figures 6, 7:
Lectotype. Figures 8-10. Holospira maxwelli Pilsbry, 1953. Paratype (UF 50207). Scale bar a is for Figures 6, 7;
scale bar b is for Figures 8, 9; scale bar c is for Figure 10.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 11-16. Holospira acanthidia n. sp. Figures 11, 12: Holotype (UF34377). Figures 13-16:
Paratypes (UF 80952). Scale bar a for Figures 11-15. Scale bar b for Figure 16.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Nr. '


I~


Figures 17-24. Holospira rehderi Bartsch, 1947. Figures 17, 18: Holotype (USNM 543590). Figures 19, 20:
paratypes (USNM 543590). Figure 21: Holotype of Holospira morelosensis Bartsch, 1947 (USNM 543591).
Figures 22-24: paratypes of Holospira morelosensis (USNM 543592). Scale bar for Figures 17-24.


I jlr"
-40ii~i~


''* *





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 25-31. Holospira rehderi Bartsch, 1947. Morelos, 2 km NW of Nopalera (UF
200399). Scale bar a for Figures 25-30. Scale bar b for Figure 31.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 32-39. Holospira rehderi Bartsch, 1947. Figures 32-35: Morelos, 7 km W of Jojutla (UF 34287). Figures
36-39: Puebla, 13 km SE of Izucar de Matamoros (UF 34373). Scale bar for Figures 32-39.


Gc-





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 40-44. Holospira albertoi Bartsch, 1947. Figures 40, 41: Holotype (USNM 543495). Figures 42-44:
Puebla, 5 km NW ofPetlalcingo (UF 34383). Scale bar for Figures 40-44.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 45-52. Holospira zygoptyx n. sp. Figure 45: Holotype (UF190771). Figures 46-52: paratypes (UF
337051). Scale bar a for Figures 45-49, 52. Scale bar b for Figures 50, 51.


tj





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 53-59. Holospira. Figures 53-56: Holospira melea Bartsch, 1926. Puebla, Ixcaquixtla, Holotype (USNM
363145). Figures 57-50: Holospira hyperia Bartsch, 1926. Puebla, Esperanza, Holotype (USNM 363146). Scale
bar for Figures 53-59.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


4 -


Figures 60-66. Holospira eburnea n. sp. Puebla, limestone hilltop 7.6 km SSW of Molcaxac. Figure 61:
Holotype (UF 34294). Figures 60, 62-66: Paratypes (UF 351576). Scale bar for Figures 60-66.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


w


73 74


Figures 67-75. Holospira teres (Menke, 1847). Figures 67-70: Lectotype (SMF 7023). Figures 71-75:
Lectoparatype (BMNH 1996135). Scale bar for Figures 67-75.


,eS


N'- J'I




BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


A


p4-
77 078 79 80





















Figures 76-85. Holospira hogeana Martens, 1897. Figures 76-81: Lectotype, Veracruz, Maltrata (BMNH
1901.6.22.1903). Figures 82-85: Puebla, 1 km NE of Chapulco (UF 34376). Scale bar a for Figures 76-84. Scale bar
b for Figure 85.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 86-100. Holospira teotitlana Bartsch, 1945. Figures 86-90: Oaxaca, 2.5 km N of San Juan de Los Cues
(UF 337284). Figures 91-95: Oaxaca, 8 km S of Teotilan de Camino (UF 200425). Figures 96-100: Puebla, 11.5 km
SSE ofCoxcatlan (UF 337317). Scale bar for Figures 86-89, 91-100.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


1 102 103 104 105




























Figures 101-109. Holospira. Figures 101-105: Holospira pfeifferi (Menke, 1847), Puebla, Tehuacan; Lectotype
(BMNH 1996134). Figures 106-109: Holospira oaxacana Bartsch, 1906, Oaxaca, Tomallin. Figures 106 Lecto-
type (USNM 175085). Figures 107-109: Paralectotypes (USNM 1025749). Figure 109 is the internal barrier of
Figure 108. Scale bar is for Figures 101-108.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 110-116. Holospira rhinion n. sp. Puebla, 15 km SE of Tehuacin. Figure 110: Holotype (UF 34375).
Figures 11-116: Paratypes (UF 342782). Figure 15 is an enlargement of Figure 14. Scale bar is for Figures 110-114,
116.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


40 .x








Figures 117-122. Holospirapainteri Bartsch, 1906. Puebla, Tehucan. Figure 117: Holotype (USNM 187675).
Figures 118-122: Paratypes (USNM 1025750). Figure 122 is an enlargement of Figure 121. Scale bar is for Figures
117-121





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 123-129. Holospira denserpens n. sp. Puebla, 1 km E of Azumbilla, 23 km N of Tehuacin. Figure 124:
Holotype (UF 190783). Figures 123, 125-129: Paratypes (UF 341543). Figure 128 is an enlargement of Figure 127.
Scale bar is for Figures 123-127, 129.


*J;Y





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 130-136. Holospira aurantiaca n. sp. Puebla, canyon 2 km SE of Tecamachalco. Figure 130: Holotype
(UF 190765). Figures 131-136: Paratypes (UF 341545). Scale bar is for Figures 130-135.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 137-144. Holospira scololaema n. sp. Puebla, 1 km NE ofYehualtepec. Figure 137: Holotype (UF 34287).
Figures 138-142: Paratypes(UF 335731). Figures 143, 144: Paratype(UF 335727). Figure 144 is an enlargement of
Figure 143. Scale bar a is for Figures 137-142. Scale bar b is for Figure 143. Scale bar c is for Figure 144.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


ep


Figures 145-150. Holospira colymis n. sp. Puebla, limestone hillside 13.5 km WSW of San Bartola Teontepec.
Figur. 145: Holotype (UF 200782). Figures 146-150: Paratypes (UF 335728). Scale bar is for Figures 145-147, 149-
150.




THOMPSON and MAHALCIK: Southern Mexican holospirid snails


-V















Figures 151-158. Holospira fortisculpta n. sp. Puebla, 5 km NW of Atenco. Figure 151: Holotype (UF 34293).
Figures 152-158: Paratypes (UF 342779). Figure 156 is an enlargement of Figure 155. In Figure 157 the parietal
lamella has been broken to expose the columellar lamella. Scale bar is for Figures 151-155, 157-158.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 159-164. Holospira haploplax n. sp. Puebla, 5 km NW of Atenco, Figure 163: Holotype (UF 34293).
Figures 164-168: Paratypes (UF 342779). Figure 168 is an enlargement of Figure 167. Scale bar is for Figures 163-
167.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


'Li9.�? .-.


L fY j


Figures 165-170. Bostrichocentrum and Holospira (Stalactella). Figures. 165-168. Bostrichocentrum tryoni
(Pfeiffer, 1867). Puebla, Puebla (UF 179808). Figure 168 is an enlargement of Fig. 167. Figures 169-171. Holospira
(Stalactella) rosei Bartsch, 1906. Puebla, Tehuacin. Holotype (USNM 188181). Figure 171 is the opened poste-
rior side of Figure 169 enlarged. Scale bar is for Figures 165-167, 168-170.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 172-177. Holospira (Stalactella) cremnobates n. sp. Puebla, 10 km N of Tehuacin. Figures 172, 173,
173a: Holotype (UF 190789). Figures 174-177: Paratypes (UF 21453). Scale bar is for Figures 172-176.





THOMPSON and MAHALCIK: Southern Mexican holospirid snails


Figures 178-183. Holospira (Stalactella) psectra n. sp. Puebla, a limestone cliff on the S slope of Cerro Caolalote,
2 km E of El Carmen. Figures 178, 179: Holotype (UF 233197). Figures 180-183: Paratypes (UF 41472). Scale bar
is for Figures 178-182.




BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


Figures 184-190. Holospira (Stalactella) marmorata n. sp. Puebla, 13.5 km WSW of San Bartolo Teontepec.
Figures 184, 185: Holotype (UF 233188). Figures 186-190: Paratypes (UF 80954). Scale bar a is for Figures 184-
189. Scale bar b is for Figure 190.


^






THOMPSON and MAHALCIK: Southern Mexican holospirid snails


.
,-r \

,.


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Figures 191-195. Holospira (Stalactella) chazumbae n. sp. Oaxaca, 2 km NE of Santiago Chazumba. Figures 191,
192: Holotype (UF 34299). Figures 193-195: Paratypes (UF 214439). Scale bar a is for Figures 191-194. Scale bar
b is for Figure 195.


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,'"\





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(1)


4----


Figures 196-198. Coelostemma microstoma (Pfeiffer, 1861). Lectotype, British Museum (Natural History) 1996162.
Scale bar is for Figures 196-198.







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