New and little known land snails of the family Spiraxidae from Central America and Mexico (Gastropoda, Pulmonata)

Material Information

New and little known land snails of the family Spiraxidae from Central America and Mexico (Gastropoda, Pulmonata)
Series Title:
Bulletin - Florida Museum of Natural History ; volume 39, number 2
Running title:
Spiraxidae from Central America and Mexico
Thompson, Fred G ( Fred Gilbert ), 1934-
Place of Publication:
Gainesville, Fla.
University of Florida
Publication Date:
Copyright Date:
Physical Description:
p. 45-85 : ill. ; 23 cm.


Subjects / Keywords:
Spiraxidae -- Central America ( lcsh )
Spiraxidae -- Mexico ( lcsh )
Snails -- Central America ( lcsh )
Snails -- Mexico ( lcsh )
Florida Museum of Natural History ( local )
City of Gainesville ( local )
Holotypes ( jstor )
Paratypes ( jstor )
New species ( jstor )
bibliography ( marcgt )
non-fiction ( marcgt )
Spatial Coverage:


Includes bibliographical references (p. 84-85).
Abstract also in Spanish.
General Note:
Cover title.
Statement of Responsibility:
Fred G. Thompson.

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University of Florida
Holding Location:
University of Florida
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Copyright held by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, and personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum. For permission or additional information, please contact the current editor of the Bulletin at
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Fred G. Thompson

Biological Sciences, Volume 39, Number 2, pp. 45-85



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Publication date: November 6, 1995

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Fred G. Thompson'


Two new genera and twelve new species are described, and various taxonomic changes are proposed.
EUGLANDININAE.- Euglandina, subgen. Varicoturris Pilsbry, 1907: the subgenus is redefined.
Gheisbreghtia Baker, 1941 is a synonym. The type species, Spiraxis dubia Pfeiffer, 1856 is redescribed
and the type specimen is refigured. The subgenus is known from the Mexican states of Veracruz and
Chiapas, and from Depto. Huehuetenango, Guatemala. Two new species are described: E. (V.) pycnoptyx,
n. sp. from Veracruz, and E. (V) huehuetenangoensis, n. sp. from Guatemala.
Euglandina, subgen. Guillarmodia Baker, 1941: the subgenus is redefined. The type species, E. (G.)
pupa Baker, 1941, is figured. The subgenus is known from lowland areas in eastern and western M6xico
and contains nine species including six newly described: E. (G.) brachystyla, n. sp., E. (G.) comma, n. sp.,
E. (G.) gracilior, n. sp., E. (G.) stenotrema, n. sp. and E. (G.) arthritica, n. sp. from Guerrero and E. (G.)
king, n. sp. from Oaxaca.
Euglandina, subgenus Varicoglandina Pilsbry, 1908: E. (V.) rubiginosa, n. sp. is described from
Depto. Huehuetanango, Guatemala.
STREPTOSTYLINAE.-Myxastyla n. gen.; type species: Streptostyla coxeni Richards, 1938. Referred
species: Myxastyla pycnota n. sp., Myxastyla hyalina n. sp. The genus occurs in north-central Guatemala.,
Belize, and Honduras.
SPIRAXINAE.- Mayaxis n. gen., type species: Mayaxis leei n. sp. from Honduras. Mayaxis is known
from Chiapas, Guatemala, Belize, and Honduras. Referred species include: Achatina lirifera Morelet,
1851; Achatina chiapensis Pfeiffer, 1856; Pseudosubulina fortis Martens, 1898; Pseudosubulina
mitescens Martens, 1898; and Pseudosubulina martensi Pilsbry, 1919.


Se described dos nuevos g6neros y doce nuevas species, y se proponen various cambios taxon6micos.
EUGLANDININAE.- Euglandina, subg6n. Varicoturris Pilsbry, 1907: el subg6nero es redefinido.
Gheisbreghtia Baker, 1941 es un sin6nimo. La especie tipo, Spiraxis dubia Pfeiffer, 1856 es redescrita y el
especimen tipo es redibujado. El subg6nero es conocido en los estados mexicanos de Veracruz y Chiapas, y
en el Depto. Huehuetenago, Guatemala. Dos nuevas species son descritas: E. (V.) pycnoptyx, esp. n. en
Veracruz, yE. (V) huehuetenangoensis, esp. n. en Guatemala.

The author is Curator of Mallogy Florida Museum ofNatural History. University of Florida, P.O. Box 117800, Gainesville FL 32611-
7800, U.SA.

THOMPSON, F.G. 1995. New and little known land snails of the family Spiraxidae from Central America
and Mexico (Gastropoda, Pulmonata). Bull. Florida Mus. Nat. Hist. 39(2):45-89.


Euglandina, subg&n. Guillarmodia Baker, 1941: el subg6nero es redefmido. La especie tipo, E. (G.)
pupa Baker, 1941, es dibujada. El subgenero es conocido en las tierras bajas al este y oeste de M6xico y
consta de nueve species incluyendo seis nuevas descritas: E. (G.) brachystyla, esp. n., E. (G.) comma, esp.
n., E. (G.) gracilior, esp. n., E. (G.) stenotrema, esp. n. y E. (G.) arthritica, esp. n. en Guerrero y E. (G.)
kingi, esp. n. en Oaxaca.
Euglandina, subgenero Varicoglandina Pilsbry, 1908: E. (V.) rubiginosa, esp. n. es descrito en el
Depto. Huehuetanango, Guatemala.
STREPTOSTYLINAE.- Myxastyla gn. n.; especie tipo: Streptostyla coxeni Richards, 1938.
Species asignadas: Myxastyla pycnota esp. n., Myxastyla hyalina esp. n. El g6nero existe al norte-centro
de Guatemala, Belice y Honduras.
SPIRAXINAE.-Mayaxis n. g6n., especie tipo: Mayaxis leei esp. n. en Honduras. Mayaxis es conocido
en Chiapas, Guatemala, Belice y Honduras. Las species asignadas incluyen: Achatina lirifera Morelet,
1851; Achatina chiapensis Pfeiffer, 1856; Pseudosubulina fortis Martens, 1898; Pseudosubulina
mitescens Martens, 1898; yPseudosubulina martensi Pilsbry, 1919.


SPIRAXIDAE are carnivorous land snails similar in appearance but not closely
related to the West Indian OLEACINIDAE. SPIRAXIDAE show relationship to
ACHATINOIDEA; OLEACINIDAE are related to SAGDIDAE in the superfamily
OLEACINOIDEA (Baker, 1962). The family SPIRAXIDAE is abundantly
represented in northwestern Central America and Mexico. This large geographic
area has been very poorly explored for land snails, particularly for the obscure
smaller species that live in rock piles and in leaf-litter. Many museum collections
contain specimens of the larger and more conspicuous species of the spiraxid
genera Euglandina and Streptostyla. Most specimens with reliable data were
collected by herpetologists and mammologists. Smaller species are seldom
SPIRAXIDAE are very much understudied taxonomically because of three
major difficulties. The primary problem is that many species described in the 19th
century were poorly diagnosed or were inadequately illustrated, rendering literature
comparisons very difficult. A second problem is that subtle differences in shell
sculpture and aperture morphology are important for distinguishing closely related
species. Information depicting these subtle features frequently was omitted in
earlier descriptions. Third, in some instances generic and subgeneric divisions
were based on anatomical characteristics that are not supported by clear-cut shell
differences, rendering generic identifications based only on shells doubtful in some
instances, particularly in the subfamily Spiraxinae. These three problems cause
great difficulty and frustration to those who first attempt identifications.
During the past 38 years I have collected numerous specimens of spiraxids
from M6xico south to Bolivia. It is apparent from the material I have examined
that most species are locally endemic and have biogeographic patterns similar to
those found in the neotropical AMPHIBIA as listed in Frost (1985). The greatest
faunal diversity for the family occurs in M6xico and northern Guatemala. At some
localities in Veracruz, Chiapas, and Guatemala I have collected as many as 20
different species sympatrically. I record some of these here, as well as species from


other localities. Most of the smaller species described in this account were
recovered from leaf-litter samples and were seldom noticed in the field.
Standard measurements are used in this study. Larger specimens were
measured with Vernier calipers. Smaller specimens were measured with an ocular
micrometer in a WILD M-5 dissecting microscope. Micrometer units were
converted to metric units. They are reported to three significant places, although
this actually exceeds the accuracy of the methodology employed. It is appropriate
to elaborate on a few of these because of the distorted aperture that some species
possess. Abbreviations in parentheses are used in tables of measurements. The
height of the aperture (ApH) is measured parallel to the axis of the shell from the
outer edge of the basal lip to the outer edge of the posterior corer. The length of
the aperture (ApL) is the longest axis measured from the outside of the posterior
corer to the outside of the basal-columellar lip. The width of the aperture (ApW)
is the greatest span within the aperture perpendicular to the aperture length.
Geographic coordinates given for specimens collected after 1990 were taken in
the field with GPS receivers. Coordinates given for earlier localities were taken
from topographic maps.


I wish to thank the following people who have assisted me with this study. Peter B. Mordan (British
Museum of Natural History) kindly loaned me type specimens in his charge. I was assisted in the field by
Steven P. Christman and Elizabeth L Raiser (Florida Museum of Natural History), Eric N. Smith (University
of Texas, Arlington), Harry G. Lee, William W. Buckingham, and Edward W. Cavin (all Jacksonville,
Florida), and Eric Fernandez (San Pedro Sula, Honduras). Wendy B. Zomlefer illustrated the shells depicted
in Figs.1-25, 38-40 and 46-48. Barbara Harmon provided the illustrations for Figs. 41-45. Photographs
comprising Figs. 27-37, 54-56 were provided by W. Stanley Blomeley. Eric Fernandez (San Pedro Sula)
and Mario Espinal (Tegucigalpa) provided invaluable assistance in Honduras. Kurt Auffenberg assisted
with the final preparation of the illustrations and many other tasks that contributed to this document
Kenneth C. Emberton reviewed the manuscript and provided many useful suggestions. I am grateful to each
for their kind assistance.

Family SPIRAXIDAE Baker, 1939
Subfamily EUGLANDININAE Baker, 1941

Genus Euglandina Crosse and Fischer, 1870
Subgenus Varicoturris Pilsbry, 1907

Type species: Spiraxis dubia Pfeiffer, 1856, by original designation.
Synonym: Gheisbreghtia Baker, 1941 (type species: Euglandina flammulata
Baker, 1941).
The definition of the subgenus is amended as follows. Shell small, about 7-10
mm long; oblong-turrite, with a long spire composed of about 8-9 narrow, convex
whorls. Aperture short and diagonal; columellar plate oblique, strongly twisted
and truncate. Embryonic whorls smooth; subsequent whorls with vertical ribs or


growth threads which are enlarged at the top and form an angulate shelf or cord
below the suture; spire with occasional reddish flames that precede growth varices.
Baker (1941) differentiated Gheisbreghtia from Varicoturris as having a
subsutural cord and a more oblique columella. Various species from Mexico and
Guatemala described below are intermediate in character and obviate the
distinction of more than one subgeneric group. Another subgenus, Streptostylella
Pilsbry, 1907, is monotypic and is known only from shells. It differs by having
strongly carinate whorls sculptured with widely spaced, heavy vertical ribs.
Characteristics of the aperture are very similar to Varicoturris. Baker (1941)
treated Gheisbreghtia and Streptostylella as sections (= species groups) within
Varicoturris, which differ only by shell morphology. Streptostylella has page
priority over Varicoturris and is the senior name available for the subgenus in
event that they prove to be synonyms.
Varicoturris is known from the Mexican states of Veracruz and Chiapas, and
from the Depto. Huehuetenango, Guatemala. The subgenus includes four species.

Euglandina dubia (Pfeiffer, 1856)
Figures 1-2, 5

Spiraxis dubia Pfeiffer, 1856. Proceedings of the Zoological Society of London:
378. (Type locality: "Chiapa").- 1859. Monographia heliceorum viventium, IV:
Streptostyla dubia (Pfeiffer), Fischer and Crosse, 1875. Miss. Sci. Mex. Amer.
Cent. : 65.- Martens, 1891. Biol. Cent.-Amer., I: 90; pl. 5, fig. 10.- Pilsbry,
1907. Man. Conch.: II, 19: 161-162; pl. 28, fig. 65.
Euglandina dubia (Pfeiffer), Baker, 1941. Nautilus, 55: 54.
Until now this species has been known only from a single specimen which
came from an unspecified locality in "Chiapas" collected by M. Gheisbreght. The
figures given in Martens (1891) and in Pilsbry (1907) do not correctly depict the
species. Through the kindness of Dr. Peter B. Mordan I have had the opportunity
to examine the syntype (BMNH 1991144). It is consistent with figures published
in the works cited above, except that it has a perforation through the back of the
last half whorl. It is 8.9 mm long, contains 8.5 whorls, and has a blunt outer lip,
indicating maturity. The columellar plate is nearly vertical and is only slightly
twisted, in contrast to the much more twisted condition that normally exists at
maturity. About a quarter turn behind the aperture there is a strong and irregularly
developed varix at the base, indicating an injury at this stage of growth which
caused the development of an atypical columella.
A specimen from Tabasco (Figs. 1-2, 5) is similar to the holotype in most
respects, except that it is slightly smaller but mature with an oblique and truncate
columellar plate. The shell is shiny and consists of 8.0 whorls. The color is
tawny-gray with occasional irregularly-spaced rusty flames. The protoconch has
2.0 smooth whorls. The first whorl is raised and pointed. The teleoconch has


strong, close, obtuse plica that are raised and truncate below the suture forming a
narrow shelf-like ledge (Fig. 5); the plica diminish below the middle of the last
whorl. The aperture is short and oblique. The columella is slightly oblique in
frontal view, and its base projects forward in lateral view (Fig. 2); its base is
twisted-truncate and is thickened at its terminal edge. The outer lip is impressed
and strongly arched forward near the middle, where it is blunt-edged and thickened
internally. Shell length 8.50 mm; shell width 3.88 mm; aperture length 3.30 mm;
aperture height 3.08 mm; aperture width 1.25 mm.
Anatomy.- A single immature specimen from near IxhuatAn, Chiapas was
examined. It contained 8 whorls and was 6.5 mm long. The anatomy does not
differ conspicuously from that described by Baker (1943) for E. flammulata.
Salient features are as follow. Mantle collar with subequally developed dorsal and
ventral lappets. Mantle spotted with black dots; minor veins distinct along
anterior third of mantle. Right eye muscle free from genitalia. Penis retractor
muscle attaching to inner surface of lung. Genital atrium long and stout. Penis
and vagina underdeveloped due to immaturity. Epiphallic section of vas deferens
long and enlarged; free segment of vas deferens very short. Prostate about half as
long as the uterus. Ovotestis not seen.
Specimens Examined.- MEXICO. CHIAPAS: 3 km N of IxhuatAn, 550 m alt.
(UF 193038, specimen dissected). TABASCO: Limestone hills 4 km E of Teapa
(UF 77996).

Euglandinaflammulata Baker, 1941
Figures 3-4, 6-7

Euglandina (Gheisbreghtia) flammulata Baker, 1941. Nautilus, 55: 56-57; pl. 5,
figs. 10-11 (shell).-- Baker, 1943. Proc. Acad. Nat. Sci. Phila., 95: 7; pl. 1,
figs. 10-11 (anatomy). Type locality: Las Tortolas, nr. Cordoba, 2700 ft. alt.
A specimen is illustrated for the purpose of comparison with related species
(Figs. 3-4, 6-7).
Specimens Examined.-- MXICO. VERACRUZ: Cerro de las Palmas, 1 km E
of Berlin, ca. 4 Km N of Cordoba, 980 m alt. (UF 190813, 190855).

Euglandina pycnoptyx, new species
Figures 8-13

Shell.- Minute, adults 5.5-6.7 (5.6) mm long. Shell slender, 0.39-0.48 times
as wide as long; turrite in shape with a weakly concave spire; spire 0.61-0.66
(0.63) times length of shell. Solid, translucent; color light gray with faint and
irregularly spaced rusty flames. Whorls 7.4-8.1 (7.4), separated by an impressed
angulate suture that forms a narrow shelf and is bounded below by a weakly beaded
spiral cord. Protoconch not distinctly demarcated from teleoconch. First whorl


Figures 1-2. Euglandina dubia (Pfeiffer, 1851), (UF 77796).



," e'i '

\ ii u i
', // \ l/ ,,' ( r,; "- i__I_7

3 4 6 7

Figures 3-4,. 6-7. Euglandina flammulata Baker, 1941. (UF 190855). Fig. 5. Euglandina dubia
(Pfeiffer, 1851) Sculpture along penultimate whorl (UF 77796). Figures 6-7. Euglandina flammulata
Baker, 1941; Sculpture along penultimate whorl (UF 190855).


pointed and protruding. First two whorls smooth; following whorl gradually
acquiring faint vertical striations; remaining whorls with strong, regular vertical
plicate striations; their upper ends cutting spiral subsutural cord into a series of
irregular elongate beads (Figs. 12-13). Plicate striations interspersed with very
weak minor striations; both becoming obsolete near the base of the last whorl.
Growth varices sparse and inconspicuous. Aperture small and narrowly elongate
with a deep sub-columellar embayment; outer lip slightly advanced near middle
(Fig. 9); basal lip receded; both outer and basal lip considerably thickened within.
Columella strongly oblique and projecting forward at the base. Columellar plate
strongly twisted and thickened along edge.
The holotype is slightly immature. It was live-collected and the color pattern is
well preserved. Measurements in mm (converted from micrometer units) of the
holotype and two paratypes are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 5.56 2.50 2.06 2.31 1.00 7.4
UF 81795 6.19 2.63 2.19 2.38 1.13 7.8
UF 81914 6.69 2.63 2.25 2.50 1.25 8.1

Type Locality.- Limestone bluff 6 km NE of Comalapa, Veracruz, MEXICO
(18042'N, 96051'W); 220 m alt.. HOLOTYPE: UF 81946; collected 14 January,
1986 by Fred G. Thompson and Steven P. Christman. PARATYPES: UF 81795
(1), 81886 (2), 81898 (1), 81914 (1), 81984 (2); all from 1-6 km NE of Comalapa;
220-450 m. alt.
The area of the type locality is in a zone consisting of mesic forest over a
strongly karsted limestone substrate. Specimens were found in leaf litter.
Distribution.- Known only from the type locality.
Remarks.- This species is most similar to Euglandina flammulata Baker,
1941 (Figs. 3-4). The latter differs by having a thinner shell with a thinner
columellar fold, a more attenuate and weakly convex spire, weaker growth
striations, and a well developed spiral cord below the angulate suture (Figs. 6-7 ).
The spiral cord is not broken into beads as it is in E. pycnoptyx. The two species,
though closely related, are disjunct in distribution and do not overlap in shell
Etymology.- The species name pycnoptyx is from the Classical Greek xuiKVw
q, thickened, and Irzruoq, a fold. The name alludes to the thickened columellar
fold that characterizes this species.



I 1

Figures 8-13. Euglandinapycnoptyx, new species. Figures 8-9. Holotype (UF 81946). Figures 10-11.
Paratype (UF 81795). Figures 12-13. Holotype; sculpture along shoulder of penultimate whorl.


Euglandina huehuetenangoensis, new species
Figures 14-16

Shell.- Translucent and moderately sized for typical Gheisbreghtia, about 7.7
mm long; elongate-ovate in shape, 0.48 times as wide as long. Spire convex in
outline, 0.61 times length of shell. Tawny gray in color with frequent reddish
flames behind growth varices. Whorls 7.2.; first whorl protruding, rounded.
Protoconch with 3 smooth turgid whorls separated by a moderately impressed
suture. Teleoconch with weak, wide plicate striations that become obsolete near
the middle of the whorl. Tops of plicae forming a narrow angulate shelf at the
suture (Fig. 16), but a subsutural cord in not developed. Aperture oblique and
narrow; elongate-auriculate in shape with deep columellar and sub-columellar
embayments. Peristome thin; outer lip nearly vertical; basal lip receded.
Columella short; oblique and protruding forward (Fig. 15); columellar plate
moderately thin.
Measurements in mm (converted from micrometer units) of the holotype are as
follow, length 7.69; width 3.69; aperture height 3.00; aperture length 3.50;
aperture width 1.44.
Type Locality.- Finca Chiblac, ca. 5 km W of San Ram6n, Dept.
Huehuetenango, GUATEMALA (15052'38"N, 9114'34"N); 1000 m alt.
HOLOTYPE: UF 190305; collected 4 March, 1991 by Fred G. Thompson and
Steven P. Christman. PARATYPES: UF 190344 (2 immature specimens); cave
below Finca Chiblac, Dept. Huehuetenango, Guatemala; 700 m alt.
Distribution.- Known only from the type locality.
Remarks.- This species is unique within the subgenus because of its relatively
obese shape. It differs from Euglandina dubia and bears some similarity to E.
flammulata and E. pycnoptyx from Veracruz because of the weak sculpture. It
differs from the latter two species by having even weaker sculpture that becomes
obsolete above the middle of the whorl, by lacking any trace of a subsutural cord,
by being larger and stockier, and by having fewer whorls. Characteristics of the
suture indicate a closer relationship with E. dubia, which has a larger and more
attenuate shell and a strongly plicate sculpture that forms close round ribs.
Etymology.- The species name alludes to the province in Guatemala from
where it was collected.

Subgenus Guillarmodia Baker, 1941

Previously three species were recognized in this group: E. pupa Baker, 1941b,
E. elegans (Martens, 1895), and E. dorsalis Thompson, 1963. Guillarmodia is
redefined to include some additional and larger species. The shell is solid and
slender and is up to 26 mm long. It is sculptured with weak growth striations and
is nearly uniform in color. It lacks color bands or flames.


Figures 14-16. Euglandina huehueteangoensis, new species. Figures 14-15> Holotype (UF 190305).
Fig. 16. Sculpture along shoulder of penultimate whorl.


Guillarmodia includes nine species and is known from Veracruz, Oaxaca,
Guerrero, Michoacan, and Colima. Three species-groups are recognized. The
typical group (E. pupa) is monotypic and is confined to southern Veracruz. It has
a small elongate shell 6.75-8.13 mm long and a simple suture. A second group
consists of species that bear a raised callus-like ridge on the face of the lower
parietal wall and columella and have a simple or beveled suture. Species vary in
size from 6-18 mm long. It includes four species known from coastal zones of
Colima south to Guerrero. Another species described below from Oaxaca lacks a
callus but also may belong in this group. The third group is confined to
intermediate elevations in central Guerrero. It is characterized by its extremely
elongate-elliptical shape, larger size (15-26 mm), and complex suture along the

Euglandina pupa Baker, 1941
Figures 17-19

Euglandina (Guillarmodia) pupa Baker, 1941. Nautilus, 55:57; pl. 5, figs. 8-9
(shell).- Baker, 1943. Proc. Acad. Nat. Sci. Phila., 95: 8; pl. 2, figs 14-15
Type Locality.- AtoyAc, Veracruz, MEXICO; 1300-1415' alt. Previously this
species was known only from the type locality. A specimen from a nearby locality.
is illustrated (Figs. 17-19) for comparison with related species. The following
records add considerably to its known distribution.
Specimens Examines.- MXICO. OAXACA: limestone range 1 km W of
Cedral, 100 m alt. (UF 159530, UF 159500); 4 km NW of Temascal, 100 m alt.
(UF 159469); 3 km S of Acatlan, 100 m alt. (UF 189740); 4 km SW of Acatlan,
100 m alt. (UF 159530). VERACRUZ: 3 km NE of AtoyAc, 640 m alt. (UF
190901); 1 km NW of Atoyaquillas, ca. 7 km NNW of Paraje Nuevo, 750 m alt.
(UF 190924); limestone knoll 4 km ESE of Cordoba, 800 m alt. (UF 159365); 1
km E of Berlin, ca. 4 Km N of Cordoba, 980 m alt. (UF 190820); Cerro de Las
Palmas, 1 km N of San Mateo, ca. 3 km NE of Cordoba, 910 m alt. (UF 190874);
Comalapa, Cueva del Tunel (UF 77504, UF 77526); 4 km NE of Comalapa, 400 m
alt. (UF 81909); 6 km NE of Comalapa, 250 m alt. (UF 81945).

Euglandina brachystyla, new species
Figures 20-22

Shell.- Shell minute, 6.4-6.9 mm long; 0.42-0.44 times as wide as high.
Glossy and transparent; amber colored with a lighter-tinged columellar crest.
Elliptical in shape; apex obtuse. Spire 0.56-0.60 times length of shell; convex in
outline. Whorls 6.5-7.0; moderately arched between sutures; not shouldered; with
a clear subsutural hyaline zone. Suture simple and moderately impressed.
Protoconch weakly demarcated, consisting of about 2.5 smooth whorls.



17 18

Figures 17-19. Euglandina pupa Baker, 1941. (UF 159500). Fig. 19. Sculpture of penultimate whorl.

Teleoconch with occasional barely distinct growth striations. Aperture narrow,
height equal to or less than the width of the shell; about 0.29-0.31 times as wide as
long; constricted below the periphery of the outer lip and with a distinct
embayment at parietal-columellar corner. Columella slightly oblique, weakly
twisted and very short (Figs. 21-22). Peristome conspicuously thickened along
lower outer lip and slightly less so along the base. Parietal wall thickened and
forming a rounded crest that continues below to the base of the columellar lip.
Measurements in mm (converted from micrometer units) are as follow.



Figures 20-22. Euglandina brachystyla, new species. Figures 20-21 Holotype (UF 193587). Fig. 22.
Paratype (UF 193588).

Specimen length width ApH ApL ApW whorls

HOLOTYPE 6.38 2.79 2.79 2.98 0.87 6.5
PARATYPE 6.94 2.85 2.79 2.98 0.93 7.0


Type Locality.- A small isolated karst limestone knoll 2 km east of Punta
Troncones, Guerrero, MEXICO (17047'36"N, 101042'43"W); 60 m altitude.
HOLOTYPE: UF 193587; collected 3 March, 1992 by Fred G. Thompson.
PARATYPE: UF 193588; same data as the holotype.
Punta Troncones is a small fishing village on the Pacific Coast about 12 km
northwest of Panda, Guerrero. The type locality is at the southeastern base of an
isolated low mountain range that lies along the coast. The range consists of
igneous, metamorphic, and sedimentary formations, including a few very localized
karsted limestone outcrops. The two specimens comprising the type series were
recovered from a leaf-litter sample taken from under large boulders at the top of
the knoll. The vicinity is in a stand of dense lowland mesic deciduous forest. The
knoll is sparsely vegetated with trees and vines and bears very little understory.
Distribution.- Known only from the type locality.
Remarks.- This and E. pycnoptyx are the smallest known species of
Euglandina, a genus that varies in size up to 112 mm long (E. titan Thompson,
1987). Euglandina brachystyla is related to a group of species occurring in
southwestern Mexico that include E. elegans (Martens, 1895) E. dorsalis
Thompson, 1963, and the following species. All are known from localities near
the Pacific coast. They are alike in bearing on the lower surface of the parietal
wall a callus-like ridge that extends downward onto the face of the columella.
Euglandina brachystyla differs from these others by its much smaller size, its
abbreviate columella, and its simple suture. No close relationship between E.
brachystyla and other species of this group is apparent.
Etymology.- The species name brachystyla is from the Classical Greek fpoax
uq, short and acuXo(, and alludes to the abbreviated columellar projection at the
base of the aperture (Figs. 21-22).

Euglandina comma, new species
Figures 23-25

Shell.- Medium-sized, adults about 13-18 mm in length. Shell light brown
and glossy with a nearly white columellar crest. Shell faintly translucent or
opaque. Elongate-ovoid in shape,; about 0.35-0.41 times as wide as long. Spire
attenuate and weakly convex in outline, 0.59-0.65 times the length of the shell.
Apex narrowly rounded, first whorl slightly protruding. Mature specimens with
7.4-8.8 weakly arched whorls. Suture impressed, forming a narrow rounded shelf.
Protoconch with 2.75 rounded whorls, which are separated by a deep suture. The
first 2.5 are smooth, and the next quarter whorl has very fine vertical threads.
Teleoconch smooth except for strongly impressed and widely spaced striations
below the suture. The striations have the appearance of a series of reversed
commas (Figs 23, 25). Striations obsolete and irregular over the rest of the surface.
Occasional weak growth varices present. Aperture elongate-auriculate in shape,
with a deep bulge into the columellar embayment; aperture about 0.33-0.41 times


as wide as long. Peristome blunt-edged but not conspicuously thickened internally.
Outer lip slightly advanced near the middle; basal lip receded (Fig. 24). Columella
rounded-truncate, short and inclined at about 450; compressed laterally and
bearing on its outer surface an opaque, thick, white callus that is continuous with
and fades into the parietal callus. The columellar crest develops in specimens with
7.4-7.7 whorls. Smaller specimens lack any indication of the structure.
Specimens from the type locality are larger than those from other places.
Specimens from near Zihuatenejo are smaller and have fewer whorls at maturity.
Aside from differences in size and whorl count all of the paratypes are remarkably
alike in their proportions and other morphological features. Measurements in mm
for the holotype and five paratypes are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 16.5 6.0 6.0 6.3 2.5 8.5
UF 193039 14.9 5.7 5.5 6.0 2.4 8.3
UF 193039 18.6 6.5 6.6 6.8 2.8 8.8
UF 193040 12.0 4.9 4.6 5.0 2.0 7.8
UF 193040 13.5 5.1 5.4 5.8 2.1 7.7
UF 193040 15.1 5.5 5.4 5.9 2.3 8.7

Type Locality.- Southeast side of the Rio Ixtapa, 6 km southeast of Pantla,
Guerrero, MEXICO (17043'N, 10140'W); 20 m alt.. Pantla is a small village on
the coastal highway 23 km NW of Zihuatanejo. The type locality is in a lowland
mesic forest overlying a substrata consisting of lateritic soils and limestone.
HOLOTYPE: UF 34663; collected 28 October, 1970 by Fred G. Thompson.
PARATYPES.- GUERRERO: Same data as the holotype (UF 193039, 4
specimens); 2 km SE of Pantla, 30 m alt (UF 34662, 5 specimens; UF 199949 -
20 specimens); 3.5 km NW of Zihuatanejo, 130 m alt. (UF 34659, 1 specimen) 4.5
km NW of Zihatanejo, 100 m alt. (UF 34581, 2 specimens; UF 193040, 17
Distribution.- This species is found along coast areas of northwestern
Guerrero, Mdxico. Other records from GUERRERO: limestone hill 3 km E of
Naranjillo (UF 211729); 2 km NE Punta Troncones (17047'36"N, 10142'43"W),
50 m. alt. (UF 200193); 1.3 km N of Playa Majahua (17047'58"N, 101044'11"W),
50 m alt. ( UF 199985, UF 200009); 10 km N of La Uni6n (18000'05"N,
101045'20"W), 150 m alt. (UF 200331); 1 km SSE of La Junta (18001'21"N,
10144'58"W), 130 m alt. (UF 200344).
Remarks.- Two other species of Guillarmodia bearing a raised callus on the
columella are E. elegans (Martens, 1895) and E. dorsalis Thompson, 1963. They
occur in Colima and MichoacAn respectively. Euglandina comma is much larger
than these other two species. It is most similar to E. dorsalis by having a truncate
suture forming a narrow spiral shelf. In the latter species the ledge is sharply


L. E
/, I '

i F--J


23 24

Figures 23-25. Euglandina comma, new species. Figures 23-25.
Sculpture along suture of penultimate whorl ofholotype.


Holotype (UF 34663). Fig. 25.

angulate at the shoulder and is bordered below by close, weak striations that
crenulate the suture. In addition, the spire is relatively shorter, being about 0.55-
0.56 times the length of the shell, and it is only slightly convex in outline. In E.
comma the edge of the ledge is rounded, and the sculpture is more widely spaced
and much stronger. The spire is more attenuate, being about 0.59-0.65 times the
length of the shell, and it is more convex in shape.
Etymology.- The species name comma is from the Latin comma, a
punctuation mark, in reference to the strongly impressed sculpture below the


Euglandina kingi, new species
Figures 26-28

Shell.- Shell medium sized, about 12-14 mm long in mature specimens.
Translucent; glossy; color uniform light copper-brown. Ovate-cylindrical in shape;
about 0.35-0.38 times as wide as high. Spire elongate, convex in outline; about
0.57-0.60 times the length of the shell. Apex rounded; first whorl weakly
protruding. Whorls of apex and spire rounded and separated by a moderately
impressed suture, which tends to be more deeply constricted in larger specimens.
Suture bounded below by a lighter colored narrow cord, which is broken into beads
of irregular length. Whorl 7.3-7.8 in mature specimens. Protoconch consisting of
2.7 smooth whorls that are margined by a narrow hyaline zone. Teleoconch
sculptured with very fine incremental striation and occasional growth varices. The
striation are nearly uniform in intensity over the surface of the shell. On the first
postembryonic whorl the striations weakly crenulate the suture; the crenulations
rapidly transforms into a sub-sutural beaded cord. Aperture elongate-auriculate in
shape; about 0.36-0.41 times as wide as long; with a moderate columellar bulge.
Peristome blunt-edged at maturity. Peristome in lateral view recurved between the
suture and the periphery and retracted below so that the outer lip is advanced near
the middle. Columella obliquely truncate and variably inclined with respect to the
longitudinal axis of the shell. In some specimens the columella is nearly vertical;
in others it slopes up to about 200 from the vertical axis.
Measurements in mm of the holotype and four specimens selected for variation
are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 12.9 4.8 5.5 5.6 2.3 7.3
PARATYPE 12.9 4.6 5.4 5.5 2.0 7.4
PARATYPE 13.4 4.9 5.5 5.8 2.3 7.5
UF 77788 14.1 4.9 5.5 5.9 2.3 7.8
UF 77788 14.3 5.0 5.4 5.8 2.1 7.8

Type Locality.- Among boulders in an open semi-xeric scrub forest on a
limestone hillside, on the west side of the Rio TehuAntepec 25 km NW of the Presa
Benito Juardz (dam), Oaxaca, MEXICO; 300 m alt. HOLOTYPE: UF 34691;
collected July 20, 1966 by F. Wayne King and Fred G. Thompson. PARATYPES:
UF 193056 (19); same data as the holotype.
Distribution.- This species is confined to low and intermediate semi-xeric
habitats along the Pacific coast of Oaxaca, M6xico.
Specimens Examined.- MEXICO. OAXACA: 30 km NW, 3 km NE of
Tehudntepec, 200 m alt. (UF 34687); 13.5 km NW of TehuAntepec, 150m alt. (UF


Figures 26-31. Euglandina king, new species. Fig. 27. Holotype (UF 34641). Figs. 28-29. Paratype
(UF 193056). Figs 29-31. Euglandina gracilior, new species. Holotype (UF 194115).


77788); 24 km NW, 3 km NE of TehuAntepec, 210 m alt. (UF 77781); 3.5 km NW
of Mixtequilla, 130 n alt. (UF 77768); 26 km SE of El Camar6n, 1100 m alt. (UF
77991); 10 km N of La Ventosa, 210 m alt. (UF 78003); limestone hill 10 km E of
La Ventosa, 50 m alt. (UF 211463); 7.5 km N of La Ventosa, 100 m alt. (UF
211489); 15 km ESE of Santiago Astata (15057'53"N; 95032'59"W), 100 m alt.
(UF 211509); limestone mtn. 3 km W of Santiago Astata (1559'55"N,
95042.45"W), 100 m alt.(UF 211518); 5 km E of Santiago Astata (15058'50"N,
9538'16"W), 100 m. alt. (UF 211550).
Remarks.- This species is unique within Guillarmodia because of the
irregularly beaded subsutural cord. Other species with a subsutural cord are
truncate above the cord, forming a narrow spiral ledge, and none are beaded as in
E. kingi. Its shape, size, relatively impressed suture, and fine incremental
sculpture also distinguish it from similar species.
Etymology.- I take great satisfaction in naming this species after my long-time
friend and colleague, F. Wayne King.

Euglandina gracilior, new species
Figures 29-31

Shell.- Large for the subgenus Guillarmodia; adults about 21.7-25.5 mm long.
Elongate-elliptical in shape; about 0.31-0.33 times as wide as long. Spire tapered
and convex in outline, about 0.63-0.65 times the length of the shell. Color uniform
brown. Shell translucent, nearly clear in life; glossy. Embryonic shell consisting
of about 2.3 smooth whorls that are separated by a weakly impressed suture. The
apical whorl slightly raised and pointed. Adult shells with 9.4-9.8 whorls that are
weakly arched between the moderately impressed suture. Shoulder of whorls
beveled. Postembryonic shell sculptured with regular but weak thread-riblets that
are slightly enlarged along the suture, but become indistinguishable below the
periphery. Aperture elongate-auriculate, of medium width with a moderate
embayment into the columellar margin; about 0.35-0.39 times as wide as long.
Columella nearly vertical and truncated; relatively longer and not as forward
projecting as in the following two species. Outer lip above the periphery relatively
thin. Outer and basal lip below the periphery conspicuously thickened. Outer lip
in lateral profile weakly arched forward below the middle and retracted near the
There is considerable local variation in size and relative slenderness of the
spire. The measurements given below are based on specimens with a
conspicuously thickened lip, which is the definitive stage of adult development. In
subadults the peristome is nearly uniformly thin, and the aperture may appear
narrower and without a conspicuous columellar embayment.
Measurements in rmm of the holotype and three other adult specimens are as


Specimen length width ApH ApL ApW whorls

HOLOTYPE 22.4 7.3 8.3 8.8 3.4 9.7
UF 34596 21.7 6.9 7.8 8.1 3.1 9.4
UF 194114 25.5 8.0 8.9 9.6 3.4 9.7
UF 194116 23.4 7.3 8.5 8.8 3.4 9.8

Type Locality.- An open oak forest 12 km by road southwest of Xochilapa,
GUERRERO (17048'41"N, 99042'32"W); 1700 m altitude. HOLOTYPE: UF
194115; collected 29 February, 1992 by Fred G. Thompson. The area is an
exposed limestone substrata with clusters of palms, shrubs and Agave sp. growing
among the oaks. Snails were found under large limestone boulders in areas
shaded by large trees.
Distribution.- This species is known only from intermediate altitudes along a
limestone ridge southwest of Xochilapa, Guerrero. It is confined to a submesic
zone between 1285-2200 m altitude that is dominated by sparse growths of oaks
(Quercus sp.) on limestone substrata. I have examined specimens from the
following localities.
Specimens Examined.- MEXICO. GUERRERO: 10 km SW of Xochilapa,
1700 m alt. (UF 34596); 11.5 km SW of Xochilapa, 1750 m alt. (UF 34591); 12
km by rd. SW of Xochilapa, 1700 m alt. (UF 194117); 15 km SW of Xochilapa,
2200 m. alt. (UF 34642); 10 km SSW of Mazcala (18034'03"N, 9755'39"W), 1285
m alt (UF 194116); 10 km SSW of Mazcala (17050'50"N, 99040'25"W), 1485 m
alt. (UF 194114).
Remarks.- Euglandina gracilior and the following two species form a small
group within Guillarmodia that is found within north-central Guerrero. The group
is characterized by having the basal and outer lip conspicuously thickened below
the periphery, by their slender, elongate-elliptical shapes, and by their relatively
large size compared to other Guillarmodia. Euglandina gracilior is found south
of the Rio Balsas Basin. The other two species occur north of the basin in a small
area between and west of Iguala and Taxco. Euglandina gracilior is similar to
them in most aspects. It differs by its darker color, its larger and slightly more
robust shell, and its weakly sculpted thread-riblets. Aspects of the columella and
the suture also separate it from its two more northern relatives.
Etymology.- The species name gracilior is derived from the Latin gracilis,
meaning slender.


Euglandina stenotrema, new species
Figures 32-34

Shell.- Light brown or grayish brown in color. Shell moderately thin
throughout; translucent and shiny. Shell very slender elongate-elliptical in shape,
about 16-19 mm long, about 0.31-0.37 times as wide as high. Spire attenuate and
slightly convex in outline; apex narrowly rounded. Whorls 8.3-9.3 in mature
specimens; whorls uniformly but weakly arched between the deeply impressed and
beveled suture. Protoconch with 2.75 slightly rounded whorls which are separated
by a shallow suture; first 2.5 whorls smooth; following quarter whorl with fine
vertical striations. Teleoconch sculptured with regularly spaced thread-riblets that
only slightly crenulate the suture at their upper ends; thread-riblets becoming weak
on the middle of the whorls and obsolete on the base. Interspaces with very fine
growth striations and occasional minute granules, which are absent on the thread-
riblets. Growth varices conspicuous only on last whorl. Aperture 0.28-0.33 times
as wide as long; narrowly auriculate in shape with a shallow columellar
embayment. Outer lip and basal lip blunt-edged; slightly thickened internally;
outer lip advanced at periphery; basal lip receded (Fig. 33). Columella truncate
and slightly oblique.
Measurements in mm of the holotype and five other specimens are as follow.

Specimen length width ApH ApL ApW whorls
HOLOTYPE 19.5 5.8 6.6 7.2 2.6 9.8
PARATYPE 16.0 5.3 5.8 6.1 2.0 8.3
PARATYPE 16.9 5.5 6.3 6.6 2.1 8.7
UF 193041 17.0 5.5 6.3 6.5 2.1 8.3
UF 34627 18.0 5.7 6.5 6.9 2.0 9.0
UF 34727 19.0 5.9 6.6 6.9 2.0 9.3

Type Locality.- A limestone hillside 4.5 km ENE of IxcateopAn,
GUERRERO, MEXICO; 2500 m alt. The area is forested with a dense growth of
junipers among limestone boulders. IxcateopAn (18030'N, 99047'W) is a small
village about 37 km WSW of Taxco. HOLOTYPE: UF 193042; collected 1
October, 1970 by Fred G. Thompson. PARATYPES: UF 34640: same data as the
Distribution.- This species is confined to a small area of limestone terrain in
northern Guerrero north of the Rio Balsas from 1100-2100 m altitude.
Specimens Examined.- MEXICO. GUERRERO: ca 10 km NE of Chapa,
1420 m alt. (18025'51"N, 99044'31"W) (UF 194108); 1 km ENE of IxcateopAn,
2250 m alt. (UF 34592); 2 km ENE of Ixcateopan, 2100 m alt (18030'16"N,
99046'56"W) (UF 194109); 7.5 km ENE of Ixcateopin, 1650 m alt. (UF 193041); 5
km E of Teloloapan, 1850 m alt. (UF 34589); 25 km E of Teloloapan, 1850 m alt.
(UF 34597, 34627); 18 km N of Tonalapa; 1100 m alt. (UF 77735); 9 km S of


Figures 32-37. Figs. 32-34: Euglandina stenorema, new species. Holotype (UF 34640). Fig. 34.
Sculpture along shoulder of penultimatye whorl. Figs 35-37: Euglandina arthritica, new species.
Holotype (UF 34666). Fig. 37. Sculpture along shoulder of penultimate whorl.


Buenavista de Cuellar, 1350 m alt. (UF 77998); ca. 20 km NNW of Buenavista de
Cuellar 1540 m alt. (18033'57"N, 99027'57"W) (UF 194110); Cerro Tuxpan, 1450
m alt. (18023'36"N, 99028'53"W) (UF 194107). Specimens were found in oak and
juniper forests under limestone boulders.
Remarks.- This is a medium-sized Guillarmodia, characterized by its very
slender shape, its weakly ribbed sculpture, and its very narrow aperture. It is most
similar to the preceding species because of the shape of the aperture, its large
number of whorls, and its long attenuate spire. It differs by its smaller size, its
narrower aperture, its lighter color and its coarse sculpture.
Etymology.- The name stenotrema is derived from the Classical Greek, aosv
og, narrow, and rptl Ca, small hole, and alludes to the narrow aperture
characteristic of this species.

Euglandina arthritica, new species
Figures 35-37

Shell.- Large, mature specimens 23-24 mm in length. Very elongate-elliptical
in shape; about 0.31-0.33 times as wide as long. Spire tapered and slightly convex
in outline, nearly straight sided; about 0.67-0.69 times the length of the shell.
Color chestnut brown with a whitish subsutural cord. Shell opaque and almost
lusterless. Whorls 9.2-9.3. Protoconch consisting of 2 whorls. Initially these are
sculptured with faint and irregular indentations which fade into weak threads by
the last 1/4 whorl of protoconch. Apical whorl slightly raised and obtusely
pointed. Suture between first few whorls not impressed so that apex forms a
smoothly rounded dome. Suture between lower whorls deeply impressed and
crenulate. Whorls of teleoconch weakly and evenly arched. Sculpture consisting
of close, fine ribs which are strongest and most distinct on the upper whorls and
become weaker and less discrete on the last whorl. Upper ends of ribs knob-like
and coalescing to form a rugose subsutural cord that weakly crenulates the suture.
Interspaces between riblets with weak incremental striations. Interspace and
surface of riblets bearing scattered, minute, elongate granules. Growth varices
conspicuous only on last whorl. Aperture elongate-auriculate, about 0.37-0.40
times as wide as long; moderately wide with a deep embayment into the columellar
margin. Peristome blunt-edged and slightly thickened internally along lower outer
lip and basal lip. Outer lip slightly advanced near middle; basal lip receded.
Columella truncate and oblique, lying at about 200 to axis of shell.
Measurements in mm of the holotype and two paratypes are as follow.

Specimen length width ApH ApL ApW whorls stage

HOLOTYPE 23.2 7.2 7.3 7.8 2.9 9.3 mature
PARATYPE 24.0 7.8 7.9 8.5 3.3 9.4 mature
PARATYPE 17.1 6.2 6.3 6.6 2.6 8.0 immat


Type Locality.- An oak-forested limestone hillside 7.5 km ENE of
Ixcateopin, Guerrero, MEXICO; 2650 m alt. HOLOTYPE: UF 34666; collected 1
October, 1970 by Fred G. Thompson. PARATYPES: (UF 193044); same data as
the holotype.
Distribution.- Known only from the type locality.
Remarks.- This species is most similar in appearance to Euglandina
stenotrema. It differs from the latter by its chestnut brown color with a whitish
subsutural cord, its wider aperture with a deep columellar embayment and an
oblique columella, its crenulate suture, and its coarser sculpture in which the upper
end of the riblets are enlarged into knobs and partially coalesce to form the
subsutural cords. In addition the protoconch is obtusely pointed and consists of 2
whorls. The suture between them is not impressed. The microsculpture of the
protoconch and the teleoconch differ as is noted in the descriptions of the two
Superficially E. arthritica and E. stenotrema are very similar. They were
collected together at the type locality of E. arthritica. A mature specimen of E.
strenotrema (UF 193041) collected at that locality is nearly identical in size to the
immature paratype of E. arthritica. It is listed for comparison in the table of
measurements for the former species.
Etymology.- The species name arthritica is from the Classical Greek ap0pvt
uq, and alludes to the knob-like swellings at the upper end of the riblets.

Subgenus Varicoglandina Pilsbry, 1908
Euglandina rubiginosa, new species
Figures 38-40

Shell.- Small, 7.6 mm long. Moderately thin and transparent. Elliptical-ovate
in shape, 0.43 times as wide as long. Spire nearly straight sided; 0.58 times the
length of the shell. Color shiny gray with faint rusty tints along the upper margin
of the whorls along suture and around the base to form two faint rusty zones that
occasionally are bridged by faint rust-colored flames preceding irregularly spaced
varices. Whorls 6.1 in only mature specimen examined. Protoconch with 1.5
smooth whorls. Teleoconch with irregular faint incremental striations that are
most distinct below the suture and diminish on the middle and lower sides of the
whorls; striations becoming stronger near aperture. Occasional weak growth
varices present. Suture deeply impressed, bordered below by a beveled narrow
hyaline zone (Fig. 40). Aperture moderately high, trapezoidal in shape with a
weakly concave columnar margin; widest below the center. Outer lip thin, slightly
advanced near middle; basal lip receded (Fig. 39). Columella moderately thin;
truncate; relatively long; oblique at about 160 to axis of shell and projected forward
at the base.


.i ,40

I 1mm
38 39 '

Figures 38-40. Euglandina rubiginosa, new species. Holotype (UF 190175).. Fig. 40. Sculpture along
shoulder of penultimate whorl.

Measurements in mm (converted from micrometer units) for the holotype are as
follow, length 7.63; width 3.25; aperture height 3.18; aperture length 3.50;
aperture width 1.63.
Type Locality.- A limestone knoll 11 km S of CobAn, Dept. Alta Verapaz,
GUATEMALA; 1350 m alt. (15024'57"N, 90024'09"W). HOLOTYPE: UF
190175; collected 24 February, 1991 by Fred G. Thompson and Steven P.
Distribution.- This species is known only from a small area in Alta Verapaz
Dept., Guatemala.
Specimens Examined.- GUATEMALA: DEPT. ALTA VERAPAZ; 15 km by
road N of CobAn, 1050 m alt. (15037'14"N, 90019'10"W) (UF 189913, 1
PARATYPE); limestone knoll 17.5 km NW of Tactic, 1330 m alt. (15021'29"N,


90025'25"W) (UF 189844, 2 PARATYPES); 2 km ESE of Cajaj, 1250 m alt.
(15033'25HN, 90006'56"W), (UF 190020).
Remarks.- Euglandina rubiginosa is characterized by its small size, its
elliptical-ovoid shape with a nearly straight-sided spire, its oblique, twisted
columella, its thin transparent shell and its rusty color pattern. The three paratypes
are fresh juvenile shells. They are like the holotype in these features.
The relationship of E. rubiginosa is uncertain. It is placed in Varicoglandina
because of its reduced sculpture, simple suture, and flammulate color pattern.
Baker (1941) recognized three species groups (sections), which he distinguished by
varying degrees of sculpture and color banding. The conical spire, striate
sculpture, and color pattern suggest a relationship to the typical species group of
Varicoturris (E. monilifera group) from Chiapas and Guatemala. Other species of
this group are much larger in size and are more distinctly flammulate (Pilsbry,
1908; Baker, 1941).
Euglandina rubiginosa is superficially similar in size and appearance to E.
(Guillarmodia) pupa Baker, 1941 (Figs. 17-19). The latter has a thick, elongate-
ovoid shaped shell that is uniform buff or milky white in color and is translucent. I
have examined numerous juvenile specimens of E. pupa, and all show the same
thick aspect of the shell as do the adults. E. pupa differs further by having a less
convex-sided spire with a shallower suture between the whorls, a broader
subsutural hyaline zone, and a relatively shorter, thicker columella in the aperture.
Also, the columella of E. pupa is vertical or nearly so, in contrast to the more
oblique columella in E. rubiginosa.
Etymology.- The species name rubiginosa is from the Latin, meaning rust-
colored, in reference to its color pattern.

Euglandina constricta, new species
Figures 41-43

Shell.- Moderately small in size, adults about 5.5-6.3 mm long, and about
0.56-0.57 times as wide as high. Glossy, hyaline; with sparse rusty flames on a
tan-colored background; edge of columellar plate white. Obovate in shape with a
conical apex that is about half the length of the shell. Last whorl with a weak
spiral constriction just below its middle. Whorls 5.9-6.0 in mature specimens.
Suture deeply impressed, forming a narrow rounded shelf along following whorl.
Protoconch consisting of 2.3 whorls that bear very fine, close vertical striation.
Following whorls with similar fine striations that become irregularly spaced and
form occasional very weak growth varices; shoulder of whorls with distinct
comma-like indentations that weakly crenulate the suture. Height of aperture
about 0.49-0.51 times the length of the shell; auriculate with a deep columellar
embayment, and slightly impressed along middle of outer lip. Aperture about
0.52-0.59 times as wide as high. Columella oblique; with a strongly twisted and



Figures 41-43. Euglandina constrica, new species.
(UF 200543).

Figures 41-42. Holotype (UF 210812). Fig. 43.


truncate columellar plate that is thickened along its outer edge. Middle of outer lip
strongly advance; basal lip receded.
Measurements in mm (converted from micrometer units) for the holotype (UF
210812) and two referred specimens (UF 200543) are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 6.26 3.5 3.2 3.5 1.8 5.9
UF 200543 5.52 3.2 2.7 3.1 1.6 6.0
UF 200543 5.27 3.1 2.9 3.2 1.5 5.7

Type Locality.- A limestone ridge 3 km WNW of Mazin Grande, OAXACA,
MEXICO (180 08' 01" N, 960 21' 48"W); 200 m alt. HOLOTYPE: UF 210812;
collected 30 July, 1993 by Elizabeth Raiser. The type locality is in a rain forest at
the crest of a limestone ridge. The holotype was recovered from leaf litter collected
from along the bases of limestone ledges.
Distribution- Known only from a small area in extreme northeastern Oaxaca,
Remarks.- Two specimens from 12 km NW of Bethenia, Oaxaca (UF 200543)
differ from the holotype (Figs. 41-42) in that they appear to achieve adulthood at a
smaller size, and the incised axial sculpture below the suture is less pronounced
(Fig. 43). Other aspects of the shell are typical as described above.
This species is not particularly close in its relationship to other known species.
It is placed in the subgenus Varicoturris because of its occasional varix-like growth
striations and its flammulate color pattern. Its small size, its relatively short
conical spire, its weakly constricted last whorl, its oblique columella with a
strongly twisted, truncate columellar plate, and its advanced outer lip combine to
form very distinctive features within Varicoturris.
Etymology.- The species name constricta is from the Latin, and refers to the
medial constriction in the last whorl which, in part, characterizes this species.

Subfamily STREPTOSTYLINAE Baker, 1941
Myxastyla, new genus

Type Species.- Streptostyla coxeni Richards, 1938.
Description.- Size minute, about 2-4 mm in length; cylindrical-ovate or ovate-
elliptical in shape, with the spire comprising about half the length of the shell.
About 4-5 whorls. Sculpture simple, consisting of nearly uniformly spaced
impressed growth varices. Aperture pinched inward along the outer lip. A
strongly twisted columellar plate forms a narrow, deep channel between the
parietal wall and the edge of the plate.


This genus is placed provisionally in the Subfamily STREPTOSTYLINAE. It
differs from other streptostylid genera by its sculpture. It resembles in shape some
species of Streptostyla, but mature Streptostyla are much larger, and none has
regular spaced impressed growth varices. To a lesser extent Myxastyla resembles
some Spiraxis (Subgenus Volutaxis) because of its strongly twisted columellar
plate. However, the various species of Volutaxis have raised axial sculpture, they
are much more attenuate in shape, and they have a greater number of whorls.
Myxastyla also resembles the Jamaican species of Sigmataxis
(OLEACINIDAE) in its sculpture. However, the shell of Sigmataxis is elongate in
shape as in Spiraxis. I suspect that the similarity between Myxastyla and
Sigmataxis in sculpture is due to convergence and no close relationship exists. The
species of Sigmataxis were reviewed by Pilsbry (1907). Baker (1941a) outlined the
relationship of Sigmataxis within the OLEACINIDAE.
At present, Myxastyla is known from central Guatemala in the Provinces of
Izabil, Alta Verapaz, and Huetuetenago, and from Roatin Island, Honduras.
Three species are known, genero-type and two new species described below. Other
undescribed species are before me from Belize and Honduras. At least two
additional species are present among the material that I have examined from
Guatemala, but the specimens are too few in number to allow further taxonomic
treatment. Material from Belize and Honduras will be treated elsewhere. Within
its range Myxastyla is found in tropical wet mesic or submesic forests over
limestone substrata and occurs between 400-1350 m altitude. It is commonly
found in leaf litter.
Etymology.- The name Myxastyla (f.) is derived from the Classical Greek
puoaan oil-lamp nuzzle, and ao'cuoc, a column, alluding to the shape of the
aperture base.

Myxastyla coxeni (Richards, 1938)
Figures 44-45

Streptostyla coxeni Riichards, 1938; Proceedings American Philosophical Society,
79:172; pl. III, fig. 2.
Type Locality.- HONDURAS: Bay Islands, RoatAn Island, limestone outcrops
between Coxen Hole and West End. Holotype: ANSP 170020.
Distribution.- Apparently endemic to RoatAn Island. Other species of
Myxastyla occur elsewhere in Honduras.
Specimens Examined.- HONDURAS: Bay Islands: Roatin Island: limestone
knoll on west side of Oak Ridge Harbor, 0.5 km N Oak Ridge (16023'43" N,
86021'18"W) (UF 224466); coraline limestone outcrop, E end Mangrove Bight, 6.5
km WSW Coxen Hole (16017'10"N, 86034"40"W) (UF 224570); 1.6 km E West
Bay Beach (16016'34"N, 86035'29"W) (UF 224531); limestone ridge 0.5 km E
West Bay Beach (16016'30"N, 86035'49"W) (UF 224546); E side of Half Moon
Bay, West End Village (16018'36"N, 86035'34"W) (UF224507).


44 45

Figures 44-45. Myxastyla coxeni (Richards, 1938); Honduras, RoatAn Island, limestone ridge 0.5 km E
West Bay Beach (16-16'30"N, 86-35'49"W) (UF 224546). Scale bar = I mm.

Remarks.-- Myxastyla coxeni is the largest of the three known species. It is
intermediate in stockyness compared to its congeners (Fig. 44-45) and is
considerably more attenuate in shape.

Myxastyla pycnota, new species
Figures 46-48

Shell- Minute, about 3 mm in length. Shell glossy, unicolor light gray,
relatively thick, translucent. Ovate-cylindrical in shape; about 0.35-0.38 times as
wide as long. Spire convex in outline; about 0.52-0.57 times length of shell.


Apical whorl large, with the first quarter raised and almost pointed. Whorls 4.3-
4.6; arched between the suture; with a broad subsutural hyaline zone that is about
1/4 the width of the whorls on the spire. Suture moderately impressed; rapidly
descending along the last whorl. Protoconch with 2.3 smooth whorls. Teleoconch
sculptured with nearly uniformly spaced impressed growth varices; smooth
between varices but with occasional fine longitudinal striations. Columellar plate
thin and weakly spiral within the shell (Fig. 47). In the last whorl the it rapidly
enlarges to form a thick spiral plate that emerges at the base of the aperture to form
and partially cover a narrow deep oblique channel between the parietal wall and
the plate. Aperture narrowly auriculate in shape and partially constricted near the
middle by the curvature of the outer lip. Outer lip pinched inward below the
middle, partially obstructing the aperture. Outer lip and basal lip strongly
thickened internally, and continuous in curvature and thickness with the base of
columellar plate. Peristome advanced near middle and strongly retracted below
(Fig. 45).
One paratype is a nearly fresh shell and is the basis for observations on color
and translucence. Other paratypes and the holotype are older and more opaque.
Measurements in mm of the holotype and four paratypes are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 2.95 1.02 1.30 1.36 0.62 4.5
PARATYPE 2.7 1.05 1.18 1.27 0.56 4.3
PARATYPE 2.7 0.99 1.27 1.30 0.56 4.4
PARATYPE 3.0 1.08 1.30 1.36 0.62 4.6
PARATYPE 3.1 1.12 1.36 1.40 0.62 4.6

Type Locality.- Gorge along the Rio Selequa 12 km SSE of La Democracia,
Dept. Huehuetenango, GUATEMALA; 950 m alt. HOLOTYPE: UF 190415;
collected 11 March, 1991 by Fred G. Thompson and Steven P. Christman.
PARATYPES: UF 193057 (5); same data as the holotype. Specimens were
collected in leaf litter gathered in a submesic zone along the base of a high
limestone cliff.
Distribution.- Known only from the type locality.
Remarks.- At present this species is known only from the type locality. We
collected specimens of Myxastyla resembling M. pycnota from three localities in
the Depto. Alta Verapaz. Apparently they represent one or more additional species
of Myxastyla. The three samples contain only one or two individuals, which are in
unsatisfactory condition for taxonomic treatment.
Myxastyla pycnota is readily distinguished from the following species by its
nearly cylindrical shape, its narrow aperture, its broad subsutural hyaline zone and
its thick peristome.


46 47 4E


) )' : ) I

\" \_ \\ ,,

49 0.5mm 50

Figures 46-50. Figs. 46-48: Myxastylapycnota, new species. Figures 46-47; Holotype (UF 190415). Fig.
48; Paratype (UF 193057). Figs. 48-50: Myxastyla hyalina, new species. Holotype (UF 189852).


Etymology.- The species name pycnota is from the Classical Greek
IUKvoxet, thickness, and alludes to the thickened lower part of the peristome.

Myxastyla hyalina, new species
Figures 49-50

Shell.- Very small, 3.2-3.7 mm long. Ovoid-elliptical in shape; about 0.42-
0.45 times as wide as long. Spire weakly convex, nearly straight-sided; about 0.44-
0.47 times the length of the shell. Thin, transparent, glossy; amber colored with a
whitish outer lip and columellar plate. Suture weakly impressed; rapidly
descending to the aperture along the last half whorl. Suture bordered below by a
narrow hyaline subsutural zone that is about 1/8 the width of the whorls on the
spire. Whorls 4.7-5.1 at maturity; weakly convex between the suture. Protoconch
rounded and containing 2.3 whorls. First whorl smooth and obtusely angulate with
a raised nucleus when viewed along the initial suture. The following whorl has
very fine vertical striations. Teleoconch sculptured with nearly regularly spaced
impressed growth varices that are continuous onto the base. Varices closer and
stronger near the outer lip; occasional very weak growth striations present between
the varices. Aperture irregularly auriculate in shape; pinched inward at middle by
weakly indented outer lip; aperture about 0.38-0.41 times as wide as high. Outer
lip strongly advanced near middle (Fig. 48). Basal lip retracted below. Peristome
slightly thickened along outer lip; slightly thinner below. Columellar plate about
as thick as outer lip; strongly reflected and high; forming and partially covering an
oblique narrow channel between plate and parietal wall (Fig. 47).
Measurements in mm of the holotype and paratypes are as follow.

Specimen length width ApH ApL ApW whorls

HOLOTYPE 3.7 1.6 1.86 1.92 0.74 5.1
PARATYPE 3.2 1.4 1.76 1.82 0.69 4.7
PARATYPE 3.3 1.5 1.74 1.86 0.68 4.7
PARATYPE 3.3 1.5 1.80 1.83 0.74 4.8
PARATYPE 3.3 1.5 1.80 1.86 0.68 4.9
PARATYPE 3.5 1.5 1.80 1.86 0.74 4.9

Type Locality.- Limestone knoll 17.5 km by road NW of Tactic, Dept. Alta
Verapaz, GUATEMALA (15*21'29" N, 90025'25" W); 1330 m alt. HOLOTYPE:
UF 189852; collected 16 February, 1991 by Fred G. Thompson and Steven P.
Christman. PARATYPES: UF 193058 (3); same data as the holotype. The
holotype and three paratypes are fresh shells. The area around the type locality
consisted of a dense thicket of small trees and shrubs in a lowland rain forest that
had been drastically cut over for small-crop agriculture. Specimens were found by
sifting leaf-litter.


Distribution.- Known only from Alta Verapaz Dept., Guatemala.
Specimens Examined.- GUATEMALA. DEPT. ALTA VERAPAZ: 11 km W
of San Cristobal Verapaz, 1120 m alt. (UF 193059); 1.5 km SE of San Juan
Chamelco; 1300 m alt. (UF 189875); 2 km ESE of Cajaj; 1250 m alt. (UF 190018);
11 km S of Coban; 1350 m alt. (UF 190169); 14 km N of CobAn; 990 m alt. (UF
Remarks.- This species is distinguished from Myxastyla pycnota by its
broader elliptical shape, its wider aperture, its thinner shell and its narrow
subsutural hyaline zone as well as other minor traits that are depicted in the
descriptions. It is distinguished from M. coxeni by its stockier form and relatively
larger aperture.
Etymology.-- The species name is from the Classical Greek uaXtvo;meaning
glassy in reference to the smooth, glossy shell.

Subfamily SPIRAXINAE Baker, 1939

The classification of the subfamily was reviewed by Baker (1939). He
recognized a single genus, Spiraxis C. B. Adams, 1850. This study recognizes as
genera Spiraxis C. B. Adams, 1850, Pseudosubulina Strebel and Pfeffer, 1882,
Volutaxis Stebel and Pfeffer, 1882, Rectaxis Baker, 1926, and Miraradula Baker,
1939. They are defined by Baker (1939) as subgenera within Spiraxis, because the
shell characters of some species are non-diagnostic at the generic level, even
though anatomical data justify the recognition of several genera. A new genus is
proposed for a group of large-sized species.

Mayaxis new genus
Figures 51-53

Type Species.- Mayaxis leei, new species.
Description.- A genus of the subfamily Spiraxinae characterized by having an
opaque, corneas colored, turrite-shaped shells bearing wide, nearly flat ribs that
crenulate the suture. The ribs are as wide as or wider than their interspaces. The
protoconch consists of about two narrow raised whorls, the first of which is smooth
and the second bears heavy axial ribs. The aperture has a truncate columella.
The following anatomical data are based on the type species. The radula (Fig.
53) contains a vestigial unicuspid central tooth, a single heavy bicuspid lateral, and
23 bicuspid marginals. The marginal teeth bear a very long slender entocone and a
short ectocone. Reproduction apparently is oviparous. The reproductive system
(Fig. 51) opens below the right eye stalk. The genital atrium is very short. The
penis is large and stout and has a short appendix at its apex and a larger stout
epiphallus (Fig. 52). The penis wall is thickened below the apex but lacks an
internal papilla or folds. The epiphallus has a large voluminous chamber at its
apical end and bears internally about six low longitudinal folds. The penis,



m/ / 52

Figures 51-53. Mayaxis eei, new genus & new species. Fig. 51 Reproductive system. Fig. 52 Penis and
associated structures. Fig. 53. Radular teeth; L = lateral, 5 = 4th marginal, 14 = 13 th marginal teeth


appendix, epiphallus, and descending vas deferens are enclosed within a sheath.
The penis retractor muscle is long and attaches high onto the inner wall of the
lung. The prostate is well developed. It is almost as long as and is tightly attached
to the uterus. The ovotestis (not illustrated) consists of a chain of multiple clavate
lobes located about one whorl above the albumen gland and is about one whorl
long. The primary gonoduct is slender and very thin; the secondary gonoduct is
greatly enlarged and convoluted along the inner curvature of the albumen gland.
The talon and the carrefour are imbedded within the albumen gland. The uterus is
long and voluminous. The post-uterine oviduct is about half as long as the uterus.
The vagina is vestigial. The spermatheca is appressed against the uterus at the
base of the prostate, as is usual for the subfamily, and its duct enters the female
system just above the genital atrium.
Mayaxis is unique within the SPIRAXINAE because of the presence of strong
wide ribs on the opaque shell, by the extremely long, slender cusps on the marginal
teeth of the radula, by the presence of an appendix on the penis, and by the
presence of an epiphallus. These characters are unique to Mayaxis and strongly
isolate it within the subfamily. Its radula is similar to Volutaxis, Pseudosubulina,
and Spiraxis by having a unicuspid central, and bicuspid lateral and marginal teeth
(Baker, 1939), although none of these has the extreme development of the inner
cusp on the marginal teeth that typifies Mayaxis. It is like other Mexican and
Central American generic groups in being oviparous and having a chain of
multiple clavate lobes comprising the ovotestis, and it is like Volutaxis in having a
penis sheath. It differs from other mainland genera but is similar to the Jamaican
genus Spiraxis, by having a well developed uterus that is longer than the combined
length of the post-uterine oviduct + the vagina. However, Spiraxis is viviparous
and has a single clavate lobe comprising the ovotestis. The combination of
characters suggest that Mayaxis is primitive in its relationships to Spiraxis,
Volutaxis, and Pseudosubulina. These genera form a clade distinct from that of
Miraradula and Rectaxis. The latter two genera are alike in having tricuspid
radular central teeth.
Described species of Mayaxis were assigned to Pseudosubulina by previous
authors because of the truncate columella, a character shared by the two genera.
Mayaxis is known from the Mayan realm of Honduras, Guatemala, and
southeastern Mexico. The name alludes to this origin and its shell shape (axis
caoqGr., m. a pole or axle). The following species are placed in Mayaxis because
of their sculpture and truncate columellae. Pseudosubulina robusta Martens,
1898, may also belong in this group.

Mayaxis chiapensis (Pfeiffer, 1856)

Achatina chiapensis Pfeiffer, 1856; Proc. Zool. Soc. Lond.: 379.
Subulina chiapensis (Pfeiffer), Crosse and Fischer, 1878; Miss. Sci. Mex., I: 637;
pl. 26, figs. 2.


Pseudosubulina chiapensis (Pfeiffer), Strebel, 1882; Beitr. Mex. Land- Susswas.
Conch., V: 119; pl. 7, fig. 17.- Martens, 1898; Biol. Cent.-Amer.: 303.-
Pilsbry, 1907; Man. Conch, Ser. I, 19: 3.
Type Locality.- "Chiapas," Mexico.
Distribution.- Known from northern Chiapas and immediately adjacent
Specimens Examined.- MEXICO. CHIAPAS: S of Motozintla, 1872 m alt.
(UF 155830). TABASCO: limestone knoll 3 km E of Teapa, 120 m alt. (UF

Mayaxisfortis (Martens, 1898)

Pseudosubulina fortis Martens, 1898; Biol. Cent.-Amer.: 304; pl. 17, fig. 17.-
Pilsbry, 1907; Man. Conch., Ser. I, 19: 5; pl. 5, fig. 19.
Type Locality.- Here-in restricted to El Reposo, western Guatemala; 800 ft.
alt.. This is the locality of the specimen illustrated by Martens.
Distribution.- Martens (1898) listed several localities in western Guatemala.

Mayaxis lirifera (Morelet, 1851)

Achatina lirifera Morelet, 1851; Testacea Novissima, II: 12.
Subulina lirifera (Morelet), Crosse and Fischer, 1878; Miss. Scietifique. Mex., 1:
633; pl. 25, fig. 12.
Pseudosubulina lirifera (Morelet), Martens, 1898; Biol. Cent.-Amer., 304-305; pl.
17, figs. 20.- Pilsbry, 1907; Man. Conch., Ser II, 19: 2-3; pl. 5, figs. 14-17.
Type Locality.- Woods of Petdn, near San Luis, Guatemala.
Distribution.- Known only from the type locality and Livingston, Guatemala.

Mayaxis martensiana (Pilsbry, 1919)

Pseudosubulina martensiana Pilsbry, 1919; Proceedings of the Academy of Natural
Sciences Philadelphia: 214; pl. 11, fig. 3.
Type Locality.- Mountains west of Livingston, Depto. Izabal, Guatemala.
Distribution.- Known only from the type locality and Las Escobas, Depto.
Izabal, Guatemala (UF 193388).

Mayaxis mitescens (Martens, 1898)

Pseudosubulina mitescens Martens, 1898; Biol. Cent.-Amer.: 304; pl. 17, figs. 18.-
Pilsbry, 1907; Man. Conch., Ser. II, 19: 5-6; pl. 5, fig. 10.
Type Locality.- Dueflas, near Antigua, Depto. Sacatepeque, Guatemala; 5000
ft. alt.
Distribution.- Known only from the type locality.



Figures 54-56. Myaxis le, new genus & new species. Holotype (UF 193387).


Mayaxis leei, new species
Figures 54-56

Shell.- Large, exceeding 24 mm in length; about 0.23 times as wide as long.
Opaque, shiny, color yellowish-corneas. Subcylindric-turrite in shape; apex
protruding and narrow, nearly pointed; upper third of shell elongate-conical with a
weakly convex spire; lower part of shell nearly cylindrical. Whorls 11.3, separated
by a deeply impressed and telescoped suture. First two whorls smooth. The next
half whorl is sculptured with weakly defined low broad riblets, which grade into
low but strong, broadly rounded riblets that are separated by narrower interspaces
on the subsequent whorls. The ribs are equally developed throughout the length of
shell and continue unto the base of last whorl. The upper ends of ribs are narrower
and raised, and weakly crenulate the suture. Last whorl with 64 ribs; penultimate
whorl with 61. Aperture height 0.19 times length of shell; strongly prosocline and
broadly ellipsoid in shape; columella truncate at base of aperture and slightly
oblique to axis of shell; columella-parietal region with a slight hump; callus
sparsely granular. Measurements for the holotype are: length of shell 24.2 mm;
width 5.5 mm; aperture height 4.6 mm.
Type Locality.- Cerro Santa Barbara, Finca Las Quebradas, 3 km W of
Nueva Esperanza, Depto. Santa Barbara, HONDURAS; 1280 m alt. HOLOTYPE:
UF 193387; collected October 11, 1991 by Fred G. Thompson and Harry G. Lee.
The holotype was found on the ground under a wood chip in a rain forest.
Distribution.- At present it is known only from the vicinity of the type
locality. A second specimen, badly weathered, was collected in a rainforest 4.6 km
WSW of Nueva Esperanza; 1430 m alt. (UF 193385).
Remarks.- Among described taxa this species is similar to Mayaxis fortis
(Martens, 1898) and M mitescens (Martens, 1898) because of the characteristics of
its sculpture and the development of the columella in the aperture. It differs from
both by its much larger size and its shape. M. fortis and M. mitescens are subulate-
turrite in shape, reach a length of 14-16 mm and have 9-10 whorls respectively.
Mayaxis leei is the largest known species in the subfamily SPIRAXINAE.
Etymology.- This species is named in honor of Harry G. Lee of Jacksonville,
Florida, a physician and an ardent collector of mollusks. Dr. Lee provided
generous support for field work in Honduras and his collecting efforts added
greatly to our field results.


Baker, H. B. 1939. A Revision ofSpiraxis C. B. Adams. Nautilus, 53: 8-16.
Baker, H. B. 1941a. Puerto Rican Oleacininae. Nautilus, 55: 24-30; pls. 1-2.
Baker, H. B. 1941b. Outline of American Oleacininae and new species from Mexico. Nautilus, 55: 51-61;
pl. 5.
Baker, H. B. 1943. The mainland genera of American Oleacininae. Proceedings of the Academy of
Natural Sciences of Philadelphia, 95:1-14; pls. 1-3.
Baker, H. B. 1962. Puerto Rican HOLOPOPODES. Nautilus, 75: 116-121.


Fischer, P and H. Crosse. 1870-1878. Mission Scientifique au Mexique et dans L'Am6rique Centrale:
etudes les mollusques terrestres et fluviatiles, I: 1-702.
Frost, D. R. 1985. Editor: Amphibian species of the world: a taxonomic and geographic reference. i-v, 1-
732. Allen press, Lawrence Kansas.
Martens, E. von 1890-1901. Biologia Centrali-Americana. i-xxvii, 1-705; pls. 1-44.
Martens, E. von, in H. Rolle. 1895. Beitrag zur Fauna von Mexico. Nachrebl. der Deutchen
Malakozoologischen Gesellschaft, 27: 129-131.
Morelet, A. 1851. Testacea novissima Insulae Cubae et Americae Centralis, 2: 1-29.
Pfeiffer, L 1856. Descriptions of nineteen new species of land-shells from M. H. Cuming's collection,
collected by M. Gheisbreght at Chiapa, Mexico. Proceedings of the Zoological Society of London,
24: 377-381; pl. 36.
Pilsbry, H. A. 1907-1908. Manual of Conchology. II, 19: i-xxvii, 1-366; pl. 1-52.
Richards, H. G. 1938. Land Mollusca from the island of Roatan, Honduras. Proceedings of the American
Philosophical Societ, 79: 167-178; pls. I-III.
Thompson, F. G. 1963. Two Mexican species of Guillarmodia s. s. Nautilus, 76: 95-99.
Thompson, F. G. 1987. Giant carnivorous land snails from Mexico and Central America. Bulletin of the
Florida State Museum, 30: 29-52; figs. 1-32.

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