Front Cover
 Table of Contents
 The giant species of Euglandin...
 Literature cited
 Back Cover

Group Title: Bulletin of the Florida State Museum. Biological sciences
Title: Giant carnivorous land snails from Mexico and Central America
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00095799/00001
 Material Information
Title: Giant carnivorous land snails from Mexico and Central America
Series Title: Bulletin - Florida State Museum ; volume 30, number 2
Physical Description: p. 30-52 : ill., maps ; 23 cm.
Language: English
Creator: Thompson, Fred G ( Fred Gilbert ), 1934-
Florida Museum of Natural History
Donor: unknown ( endowment )
Publisher: Florida State Museum, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1987
Copyright Date: 1987
Subject: Euglandina -- Classification   ( lcsh )
Euglandina -- Classification -- Mexico   ( lcsh )
Euglandina -- Classification -- Central America   ( lcsh )
Mollusks -- Classification   ( lcsh )
Mollusks -- Classification -- Mexico   ( lcsh )
Mollusks -- Classification -- Central America   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Spatial Coverage: Mexico
Bibliography: Includes bibliographical references (p. 43).
General Note: Summary in English and Spanish.
General Note: Cover title.
Statement of Responsibility: Fred G. Thompson.
 Record Information
Bibliographic ID: UF00095799
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 - 15998303

Table of Contents
    Front Cover
        Page 27
        Page 28
    Table of Contents
        Page 29
        Page 30
        Page 31
    The giant species of Euglandina
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
    Literature cited
        Page 43
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
    Back Cover
        Page 54
Full Text

of the
Biological Sciences
Volume 30 1987 Number 2





ENCES, are published at irregular intervals. Volumes contain about 300 pages and are not
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S. DAVID WEBB, Associate Editor
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Consultants for this issue:


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This public document was promulgated at an annual cost of $2980.00 or
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Publication date: 1 April 1987

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Abstract: The taxonomy of four known species of Euglandina (Gastropoda, Pulmonata,
Spiraxidae) is reviewed, and each is redescribed. E. vanuxemensis (Lea) includes the fol-
lowing synonyms: Achatina coulter Gray, Glandina coronata Pfeiffer, Glandina uhdeana
Martens and Glandina guttata Crosse and Fischer. E. aurata (Morelet) includes as a syn-
onym Achatina lignaria Reeve. E. sowerbyana consists of two subspecies: E. s. sowerby-
ana (Pfeiffer) and E. s. estephaniae (Strebel). E. gigantea Pilsbry is monotypic. E. gigan-
tae gabbi Pilsbry is a synonym. Euglandina pan new species and Euglandina titan new
species are described from Guatemala.

RESUMEN: Se revisa la taxonomia de cuatro species conocidas de Euglandina (Gastro-
poda, Pulmonata, Spiraxidae), y se redescribe cada una de ellas. E. vanuxemensis (Lea)
incluye los siguientes sin6nimos: Achatina coulteri Gray, Glandina coronata Pfeiffer,
Glandina uhdeana Martens y Glandina guttata Crosse y Fischer. E. aurata (Morelet)
incluye a Achatina lignaria Reeve como sin6nimo. E. sowerbyana consiste de dos subes-
pecies: E. s. sowerbyana (Pfeiffer) y E. s. estephaniae (Strebel). E. gigantae es monotip-
ica. E. gigantae gabbi Pilsbry es un sin6nimo. Se describe dos species nuevas proven-
ientes de Guatemala: Euglandina pan y Euglandina titan.

INTRODUCTION .......................................................... 29
ACKNOW LEDGMENTS .. ......... ....................................... 31
ABBREVIATIONS ...................................................... .. 31
THE GIANT SPECIES OF Euglandina ........................................ 31
Euglandina vanuxemensis (Lea) ...................................... .... 32
Euglandina sowerbyana sowerbyana (Pfeiffer) .............................. 34
Euglandina sowerbyana estephaniae (Strebel) ............................... 36
Euglandina gigantea Pilsbry .............................................. 36
Euglandina pan, new species ............................................. 38
Euglandina aurata (M orelet) .............................................. 40
Euglandina titan, new species ............................................. 42
LITERATURE C ITED ........................................................ 43


The purpose of this paper is to review the taxonomy of six species of
carnivorous land snails from Mexico and Central America belonging to
the typical species group of Euglandina. Species of this group attain a

IDr. Thompson is Curator of Malacology at the Florida State Museum, University of Florida, Museum Road, Gainesville,
FL 32611.
Thompson, F. G. 1987. Giant Carnivorous Land Snails from Mexico and Central America.
Bull. Florida State Mus., Biol. Sci. 30(2):29-52.


length of over 90 mm and are closely related to E. aurata (Morelet), the
type species of Euglandina. They are large and attractive, and because
of these qualities they were among the first land snails collected by nat-
uralists. Relatively few specimens have been available to earlier review-
ers of the genus, and considerable confusion exists in the literature con-
cerning the systematics of the group. Apparently they are closely related,
but this is not a settled question. Their shells share characteristics of
sculpture and color suggesting kinship. However, the anatomy of only
E. sowerbyana (Pfeiffer) has been described (Strebel 1887:34-35), and
specimens satisfactory for dissection are not available for the other spe-
cies. Thus, important questions concerning interspecific relationships must
remain in obeisance. This group includes the largest species of Euglan-
dina. They are exceeded in size among land snails only by some species
of Achatina, Corona, Megabulimulus, and Placostylus, all of which are
Euglandina is a very large, widely distributed genus. Over one hundred
species have been described. The genus is distributed from Texas to Bo-
livia, with one species of Mexican derivation occurring in the southeast-
ern United States. The majority of the species are found in Mexico. The
genus is difficult to work with taxonomically. Most species are known
only from a few specimens, and their shells are highly variable. All too
often original descriptions lack critical details of the sculpture and em-
bryonic shell structure, and the descriptions are scattered among nine-
teenth century journals and treatises that are available in only a few li-
braries. Important references to Euglandina are Strebel (1875), von
Martens (1890), and Pilsbry (1907). Numerous species have been de-
scribed since Pilsbry. Strebel classified species within the genus based
on characteristics of the embryonic shell. Later authors followed his sys-
tem with minor alterations. Baker (1941, 1943) proposed a classification
based on shell and soft anatomical features.
Euglandina has undergone extensive adaptive radiation. Species are
found in biomes as diverse as the Sonora Desert, thorn scrub forests,
montane pine forests, cloud forests, and tropical rain forests. Most
subgenus and species groups are found in a single type of biome and
have rather discrete geographic distributions similar to those demon-
strated by reptiles and amphibians. Habitat selection and ecological-geo-
graphical isolation appear to have been key factors involved in specia-
tion. Of the six species discussed in this paper, five are arboreal in rain
forests and confined to a single physiogeographic region, and one is ter-
restrial and found in xeric habitats in the desert plains of Puebla and
adjacent states. The majority of specimens in museums were collected
by herpetologists, whose field techniques lead them to examine epiphy-

VOL. 30 NO. 2


tic habitats, such as bromeliads, and to search trees and shrubs at night
for amphibians and reptiles. The arboreal habitat of these snails is re-
flected in the streaked color pattern of their shells. Arboreal snails usu-
ally have patterned shells, whereas terrestrial species generally are uni-
color. Most species of Euglandina are terrestrial.

A number of people have contributed to this project. The contributions of each have
been important because of the collective significance of their efforts and generosity. I am
grateful to the following persons for specimens they collected: W. Ronald Heyer and Roy
W McDiarmid (National Museum of Natural History), John B. Iverson (Earlham Col-
lege), Peter A. Meylan, Diderot Gicca, and Michael Corn (Florida State Museum), Walter
B. Miller (University of Arizona), Jonathan A. Campbell (University of Kansas), Gonzalo
Halffter (Instituto Politecnico Nacional, Mexico), and Douglas E. Robins (University of
Costa Rica). The following people loaned me specimens in their care. Ruth D. Turner
(Museum of Comparative Zoology), Alex Tompa (Museum of Zoology, University of Mich-
igan), Joseph Rosewater (National Museum of Natural History), Alan Solem (Field Mu-
seum of Natural History), and William K. Emerson (American Museum of Natural His-
tory). The photographs in this paper were processed by Laurence F. Getford, Staff
Photographer at the Florida State Museum. Kurt Auffenberg assembled the illustrations
and conducted many other minor but time consuming tasks. I am grateful to all of these
individuals for the assistance they have given me.

The following standard museum designations are used in this paper.
AMNH American Museum of Natural History, New York
FMNH Field Museum of Natural History, Chicago
MCZ Museum of Comparative Zoology, Harvard University
UF Florida State Museum, University of Florida
UMMZ Museum of Zoology, University of Michigan
USNM National Museum of Natural History, Washington


This paper discusses those Mexican and Central American species of
Euglandina that are closely related to E. aurata (Morelet), the type spe-
cies of Euglandina. Five are alike in being arboreal, in having relatively
ponderous shells with pronounced axial and spiral sculpture, a crenulate
suture, and a glossy yellow-orange periostracum with occasional darker
longitudinal shreaks. A sixth Mexican species, less closely related, is also
included in this group because of its large size and similar sculpture. It
differs by its terrestrial habitat and color pattern. It is the first species to
be discussed.


Euglandina vanuxemensis (Lea)

Figures 7-9
Achatina vanuxemensis Lea 1834:84.
Glandina coulteri Gray, in Beck 1837:78.
Achatina coronata Pfeiffer 1846:158.
Glandina uhdeana Martens 1863:540.
Glandina guttata Crosse and Fischer 1869:250.
A nearly complete synonymy of each of these names was given by
Martens (1890:54,56). Subsequent citations add no information concern-
ing their taxonomic status. The most recent revisions of the genus rec-
ognize two species (Martens 1890; Pilsbry 1907): Euglandina vanuxe-
mensis (Lea) (syn.: coronata, and subspecies guttata) and E. coulteri (Gray)
(syn.: uhdeana). The material that I have examined permits only one
taxon to be recognized (see discussion).
DESCRIPTION.--Shell medium to large in size, attaining a length of
up to 100 mm. Oblong in shape, being widest near the base at maturity;
less mature specimens may be elliptical-ovate in shape; shell about 0.42-
0.47 times as wide as high. Spire variable in shape, usually convex-coni-
cal in outline (Fig. 8); some specimens have a concave-shaped spire which
may be attenuated at definitive growth. Spire 0.34-0.47 times length of
shell. Suture moderately impressed. Whorls evenly convex between su-
tures. Body whorl widest in middle in immature specimens (Fig. 7), wid-
est near base at maturity (Fig. 8). Embryonic whorls 3.2-3.4 (Fig. 3),
dome-shaped with a very weakly impressed suture. First three embry-
onic whorls smooth; next half whorl with fine close longitudinal stria-
tions. Subsequent postembryonic whorls with close axial threads that
vary greatly in intensity. Some of these coalesce dorsally to form rela-
tively widely spaced conical white denticles that crenulate the suture (Fig.
11). Denticles not demarcated from threads by spiral striations along
their bases. Axial threads crossed by widely spaced, weakly differen-
tiated spiral striations that produce vaguely defined decussations on the
axial threads (Fig. 11). Spiral sculpture restricted to upper third of whorl.
Aperture elongate auriculate in shape; widest near base. Parietal wall
convex. Columella moderately long and strongly twisted to the right at
an angle of about 27-30 to shell axis. Outer lip extended forward below
middle; basal lip retracts slightly. Ground color tan or light ochraceous-
brown; marked with scattered blurred white spots followed by darker
brown streaks extending in direction of mouth. Spots and streaks evident
even in dead bleached shells.
Measurements in mm of four specimens selected to show variation

VOL. 30 NO. 2


length width height whorls width/length apert./length
MCZ 102961 47 21 28 6.1 0.45 0.59
MCZ 254629 58 24 31 6.1 0.41 0.53
UF 34909 62 29 41 6.8 0.47 0.66
UF 20808 100 42 55 7.5 0.42 0.55

TYPE LOCALITIES.-Achatina vanuxemensis: Mexico, without nearer
indication of locality. Glandina coulteri: no locality stated. Achatina co-
ronata: Mexico, without nearer indication of locality. Glandina uhdeana:
State of Veracruz. Glandina guttata: near Pueblo (presumably the city of
Puebla in the state of Puebla).
DISTRIBUTION (FIG. 16).-Confined to south-central Mexico in the
desert plateau of Oaxaca, Puebla, and adjacent areas of Mexico and Hi-
dalgo at altitudes of 1285-2375 m.
SPECIMENS EXAMINED.-PUEBLA: 3.6 mi SW of Chapulco (UF 34909);
8.1 mi SW Izucar de Matamoros 4800 ft alt (UF 34721); Hwy. 150, 1.5
mi SW of Veracruz-Puebla border (UF 35027); Puebla (USNM 185940,
MCZ 201215, MCZ 102961-paratype of G. guttata); Tecamachalco, 7500
ft alt
(UF 21327); 1.1 mi N of Tecamachalco, 7500 ft alt (UF 21329); Tehu-
acan (UNSM 443810, USNM 522954, USNM 162305, MCZ 254626, MCZ
10716, MCZ 137982); Tepeaca (UF 34670). OAXACA: El Punto (UF 20808);
Cuicatlan (USNM 251618); Huajuapan de Leon (USNM 591572); 8.3 mi
SE of Nochistlan, 7700 ft alt (UF 34571); Hwy. 131, 3.2 mi E of turnoff
to Nacaltepec (UF 35306); 0.7 mi W of Tlapacoyan, 5000 ft alt (UF 34651).
"CAMPECHE" (USNM 192985). "MEXICO" (USNM 116746, USNM 57079,
UMMZ 10970).
OTHER RECORDS.--Martens (1890:54,56) recorded this species for the
following localities. HIDALGO: Zimapan. MEXICO: Volcan de Mexical-
cingo. OAXACA: Oaxaca; Juquila. GUERRERO: Omilteme. VERACRUZ: Ja-
lapa. Baker (1941:60) gave the following records. MEXICO: Teotihuacan;
Martens (1890: pl. 2, fig. 2) figured the specimen from Zimapan, based
on a drawing of a shell presented to Mr. Abraham Lincoln and later
deposited in the Peel Park Museum. It differs from typical E. vanuxe-
mensis by its slender form, weak subsutural crenulation and its colora-
tion, having longitudinal reddish streaks on a white background. It is
similar to another specimen I have examined from 9.5 mi SW of Pinal de
Amoles, Queretaro (UF 21329). Martens also figured the specimen from
Omilteme. It differs in shape from other specimens I have examined,


being considerably wider and ovate in form, and its subsutural denticles
are weak and poorly defined. I do not believe the specimens in question
are E. vanuxemensis. Other records given by Martens and Baker are
geographically proximal to the distribution based on specimens I have
REMARKS.-This species is variable in shape and undergoes consid-
erable ontogenetic change. The synonyms are based on varying degree
of obesity and different growth stages. Achatina vanuxemensis Lea was
based on a half grown specimen 53 mm long with a weakly convex spire.
Glandina coulteri Gray was based on a slender specimen 90 mm long,
30 mm wide, and an aperture 57 mm long (Pfeiffer 1859:642-643). Acha-
tina coronata Pfeiffer was based on an adult 88 mm long with an obese
convex spire. Glandina uhdeana Martens was founded upon a specimen
67 mm long with a narrow attenuated spire. Glandina guttata was based
on a typical medium-sized specimen 69 mm long with a regularly conical
spire. The material that I have examined does not allow any to be rec-
ognized as distinct taxa.
The date of publication of the name vanuxemensis usually is cited as
1837, the final date of publication of Volume 5 of the Transactions of the
American Philosophical Society. However, the date of publication of the
issue in which the species is described was 1834. Its description was read
before the society in 1832. Thus vanuxemensis is the oldest available
name among five synonyms, dating in publication from 1834.
Euglandina vanuxemensis is characterized among other species by its
color pattern, which is unique within the genus. Its sculptural character-
istics differentiate it from the species discussed in this paper. E. vanux-
emensis also differs from these other species in that it is terrestrial and
lives in xeric habitats at moderate to high altitudes (about 1500-2300 m).
Specimens were found at night by John B. Iverson and me crawling on
rock piles and cactae. Baker (1941:60) reported finding specimens in plant
debris in xeric stations.

Euglandina s. sowerbyana (Pfeiffer)

Figures 17-20

Achatina sowerbyana Pfeiffer 1846:32.-Pfeiffer, 1853:292.-Reeve 1849:pl. 8, figs. 26.
Glandina sowerbyana (Pfeiffer) Albers 1850:198.-Fischer and Crosse 1870:98.-Strebel
1875:15; pl. 5, figs. 10a-11.-Strebel 1878:34-44, pls. 15-21 (anatomy).-Martens
Oleacina sowerbyana (Pfeiffer) Gray 1855:33.-Tryon 1885:36, pl. 6, fig. 86.
Euglandina sowerbyana (Pfeiffer) Pilsbry 1907:186.
Glandina lignaria (Reeve) Fischer and Crosse 1870:97; pl. 3, fig. 1.

VOL. 30 NO. 2


DESCRIPTION.-Shell large, up to 110 mm in length; moderately
stocky, being about 0.42-0.48 times as wide s high. Spire 0.42-0.49
times length of shell, variable in contour; most specimens have a convex
outline (Fig. 17); some have an attenuated spire that is concave in outline
(Fig. 19), as in E. gigantea. Specimens over 85 mm long with 7.0-7.5
whorls. Suture moderately impressed with whorls evenly convex be-
tween sutures. Embryonic whorls 3; apical whorl raised and dome-shaped
(Fig. 1), not coiling in a plane as in E. gigantea. Embryonic whorls ini-
tially smooth, but third whorl with comma-like striations along suture
which continue as well-defined growth striations on subsequent whorls.
Postembryonic whorls with nearly uniform, moderate, incremental stria-
tions and spiral striations (Fig. 10). The latter decrease slightly in inten-
sity near base, and decussate incremental striations to form elongate
granules that are about twice as high as wide; granules strongest on up-
per half of whorls. Suture bordered by a band of elongate bead-like den-
ticles that are nearly uniform in size and are demarcated by a weak spiral
groove. Aperture elongate-auriculate, being widest near the base; about
0.51-0.58 times length of shell. Parietal wall strongly convex. Columella
short and twisted to the right at an angle of about 300 to the axis of the
shell; forming a continuous curvature with the left basal side of the body
whorl. Outer lip arched forward below periphery.
Measurements in mm of three large specimens follow.

length width height whorls width/length apert./length
MCZ 219466 85 39 54 7.0 0.44 0.64
USNM 128291 97 41 55 7.2 0.42 0.57
UF 21334 109 52 61 7.5 0.48 0.58

TYPE LOCALITY. -Totontepec, Oaxaca, Mexico. Holotype in British
Museum (Natural History).
DISTRIBUTION (FIG. 16).-Known from eastern Mexico in the states
of Veracruz and immediately adjacent areas of Oaxaca and Puebla.
SPECIMENS EXAMINED.-PUEBLA: Puerto Morales, nr. Acultzingo 2770
m. alt. (MCZ 219466. 8 specimens). VERACRUZ: Catamaco (UF 21334);
Cerro Mano Blanco, nr. Catamaco (MCZ 201236); Cerro Chicahuaxtla
nr. Cuantlapan (UF 39852); Jalapa (USNM 512042); pine forest nr. Vera-
cruz (USNM 526229); Volcan San Martin (UF 20807); Volcan Tuxla (Vol-
can San Martin), 1475 m alt (USNM 128291). "MExICO": (USNM 317426,
USNM 10529, MCZ 10707).
OTHER RECORDS.-Martens (1890:55) recorded E. s. sowerbyana from
the following localities in the state ofVERACRUZ: Misantla; San Jose; San


Juan Miachutlan; Pacho; Jalapa; Mirador; Cerro Necoxtla between Jalapa
and Orizaba, 3000-5000 ft; Orizaba.
Martens listed a specimen collected at Omilteme, Guerrero, by H.H.
Smith. This record is dubious because such a biogeographic distribution
is unprecedented among land snails. Records from Guatemala refer to
Euglandina pan Thompson. Records from Costa Rica and Panama refer
to Euglandina gigantea Pilsbry.
REMARKS.-Euglandina s. sowerbyana is a well differentiated snail
with a restricted distribution in eastern Mexico. It is most similar to E.
gigantea, but differs from the latter as is discussed under that species.
Like E. gigantea it is an arboreal snail. Strebel (1878:35) stated that live
specimens were found on trees and banana plants, as well as in vegeta-
tive debris on the ground. Roy McDiarmid collected a live specimen in
an arboreal bromeliad (UF 29852).

Euglandina sowerbyana estephaniae (Strebel)

Figures 21-22

Glandina sowerbyana form D, Strebel 1875:17-18; pl.3, figs. 3, 3a.
Glandina estephaniae Strebel 1878:45; pl. 16, figs. 1-8 (anatomy).-Tryon 1885:36; pl. 8,
fig. 5.
Euglandina sowerbyana estephaniae (Strebel) Pilsbry 1907:186.
This subspecies differs from typical sowerbyana by its smaller size and
slender form. The largest specimen I have examined is 88 mm long
(UMMZ 10969). Measurements taken from Strebel (1895:17) and the few
specimens I have examined give a width/height ratio of 0.38-0.45.
TYPE LOCALITY. -Miahuatlan, between Jalapa and Misantla, Vera-
cruz, Mexico.
DISTRIBUTION (FIG. 16).-This subspecies is known only from a small
area around the type locality. I have examined only one specimen from a
specific locality, Jalapa (FMNH 75). Baker (1941:60) collected specimens
from Cordoba to Sumidero, 2625-3400 ft altitude.

Euglandina gigantea Pilsbry

Figures 25-26

Euglandina sowerbyana form B, Strebel 1875:16; pl. 5A, figs. 10.
Euglandina gigantea Pilsbry 1926:128; pl. 11, figs. 5-7; pl. 10, fig. 8.
Euglandina gigantea gabbi Pilsbry 1926; pl. 11, figs. 1, 2.
DESCRIPTION.-Shell large, attaining a length of 105 mm; stocky, 0.44-
0.48 times as wide as high. Spire conical, and straight sided (Fig. 25) or

VOL. 30 NO. 2


concave in outline (Fig. 26), 0.40-0.48 times height of shell. Specimens
over 60 mm long with 6.0-7.3 whorls. Suture moderately impressed;
whorls evenly convex between sutures. Embryonic whorls 3.4-3.6,
smooth. Apex blunt; first whorl revolving in a plane (Fig. 2), not raised
as in E. sowerbyana (Fig. 1). Next two embryonic whorls conical with a
weakly impressed suture. Following postembryonic whorls sculptured
with strong growth striations and spiral striations (Fig. 12). Spiral stria-
tions strongest on upper half of whorls, weak near base. Spiral striations
decussating growth striations to form elongate granules that are slightly
higher than wide. Suture bordered by a band of nearly uniform-sized
elongate beads that weakly crenulate the suture. Aperture about 0.52-
0.60 times length of shell; narrowly auriculate in shape with a nearly
straight parietal wall. Columella elongate, strongly twisted to the right
to lie at an angle of 23-28 to the axis so as to form a long straight profile
with the left base of the shell (Fig. 26). Outer lip arched forward in lateral
profile. Color yellowish with darker orange longitudinal streaks irregu-
larly spaced along lines of growth. Interior of aperture with a white tinge.
Measurements in mm based on five specimens selected to show max-
imum variation follow.

length width height whorls width/length apert./length
UMMZ 48711 90 43 52 7.3 0.48 0.58
UF 20751 89 42 49 6.7 0.47 0.57
USNM 568778 81 36 42 7.3 0.44 0.57
UF 21138 72 34 43 6.0 0.47 0.60
MCZ 211237 62 28 42 6.3 0.45 0.68

TYPE LOCALITY.-Euglandina gigantea: "Salinas Bay, in southwest-
ern Costa Rica." This is amended to Bahia de Salinas, near La Cruz,
Guanacaste Provence, Costa Rica. No locality was given for E. gigantea
gabbi. The holotypes of both are in the Academy of Natural Sciences,
DISTRIBUTION (FIG. 27).-Widely distributed in Costa Rica from near
sea-level to 2600 m altitude. Also known from Chiriqui Province, Pan-
21138, 2 specimens). Cartago Prov.: Estrella, nr. Cartago, 1500 m alt
(MCZ 211237); Tapanti, 1300 m alt (UF 39851, 39854). Guanacaste Prov.:
Finca El Silencio, 850 m alt (UF 20751); Nicoya (MCZ 77614); Tilaran
(USNM 568777, 568778). Puntarenas Prov.: Coto, 10 m alt (UNSM
522573, 522574); Rincon (UF 39855); 2.5 km NE of Monte Verde (UF


39853); Terraba, 700 m alt. (USNM 190272). "COSTA RICA" (no addi-
tional data): (AMNH 40405, FMNH 36297, MCZ 1777, UNSM 365585).
PANAMA--Chiriqui Prov.: Boquete (UMMZ 48711).
OTHER RECORDS.-Martens (1890-1901:55, 601) gave the following
records for this species as E. sowerbyana. COSTA RICA-Alahuela Prov.:
San Carlos; Turubares, 250 m (Turrucares ?). Cartago Prov.: Cache (Cachi).
Guanacaste Prov.: Rio Jesus Maria. Heredia Prov.: Sarapiqui (on Rio
Sarapiqui); Puerto Viejo. Limon Prov.: Reventazon, 500 m (on Rio Re-
ventazon). San Jose Prov.: San Jose, 1161 m. PANAMA-Chiriqui Prov.:
Chiriqui; Volcan de Chiriqui, 3000 ft.
Pilsbry (1926:128) corrected earlier misidentifications for E. gigantea,
and added Bahia de Salinas (see TYPE LOCALITY).
REMARKS.-Euglandina gigantea is most similar to E. sowerbyana in
general appearance and size. It is characterized by the narrow shape of
its aperture with a nearly straight parietal wall, its long columella that is
twisted to the right to form a long straight profile with the left base of
the shell, its blunt apex, and its relatively short granular sculpture. E.
sowerbyana has a wide, tear-shaped aperture with a convex parietal wall,
the columella is shorter and forms a weakly convex curvature with the
left basal side, the apical whorl is raised and knob-like, and the granular
sculpture is elongate, being about twice as high as wide.
Aside from the shape of the spire, E. gigantea shows little variation
throughout its range. The shape of the spire does not seem to have any
geographic or altitudinal correlations. Martens (1901:610), who discussed
this species as E. sowerbyana, quoted Biolley as stating that specimens
from the elevated central planes of Costa Rica are smaller than those
from lower elevations. I have found no such correlation among the spec-
imens I have examined. The largest specimen before me was collected
at 850 m altitude (UF 20751).
Pilsbry (1926:129) differentiated E. gigantea gabbi from E. g. gigantea
by having a smaller, thinner shell and weaker sculpture. These charac-
teristics are typical for immature gigantea, and they have no taxonomic
significance, especially in light of the fact that his specimens of gabbi
came from an unspecified locality.
All specimens that I have examined for which ecological data are avail-
able indicate that this species is arboreal. It has been found among ar-
boreal ferns and bromeliads and crawling on three trunks at night.

Euglandina pan, new species

Figures 28-29
DESCRIPTION.-Shell medium-sized, attaining a length of 70 mm.

VOL. 30 NO. 2


Moderately obese and fusiform, being about 0.42-0.46 times as wide as
high, with a moderatly inflated body whorl. Spire elongate and convex
in outline, 0.44-0.53 times length of shell, but usually shorter than height
of aperture. Whorls 7.0-7.4 (7.3 in holotype). Suture moderately im-
pressed and wavy due to crenulate sculpture. Whorls weakly convex be-
twen sutures. Embryonic whorls 3.2, smooth forming a rounded apex
(Fig. 5), first whorl planular, not raised as in E. aurata. Following two
whorls broadly convex-conical and moderately convex between sutures.
Postembryonic whorls with coarse axial striations that are irregular in
intensity and form large, white, coarse denticles along suture (Fig. 13).
Denticles variable in size and unevenly distributed along suture: not de-
marcated as a subsutural band as in E. sowerbyana or E. aurata. Axial
striations crossed by finer spiral striations which form cancellate granules
that are about half as wide as high. Spiral striations most conspicuous on
shoulder of whorl; barely distinguishable near base of shell. Aperture
narrowly auriculate in shape, about 0.47-0.56 times length of shell. Col-
umella relatively short, weakly twisted to the right at an angle of about
16-180 to shell axis (Fig. 28). Color yellowish-white with darker, irregu-
larly spaced longitudinal streaks.
Measurements in mm of four specimens follow.

length width height whorls
Paratype (MCZ 2722) 61 27 34 7.0
Paratype (FMNH 72223) 64 27 30 7.4
Paratype (UMMZ 166643 65 30 36 7.2
Holotype (USNM 487385) 70 31 38 7.3

TYPE LOCALITY.-Guatemala, Departamento Guatemala, Finca Las
Delicias, near Barillas. Holotype: UNSM 487385; collected by T. Burch
October 8, 1946.
Barillas (UMMZ 166643 1 paratype); Santo Catarina, 5 km from Gua-
temala City (FMNH 72223 1 paratype); Silvada Rd., 18 km from Gua-
temala City, 6000 ft alt (FMNH 72224 1 paratype); no data (MC2 2722
- 1 paratype).
DISTRIBUTION (FIG. 32).-Known only from moderate elevations in
central and southern Departamento Guatemala, Guatemala.
REMARKS.--Euglandina pan is distinguished from other species by its
coarse, irregular denticulate sculpture along the suture, which are not


demarked as a subsutural band, as in related species. The columella is
twisted to the right to about the same extent as in E. gigantea Pilsbry,
but it is shorter, imparting upon the aperture an auriculate shape as op-
posed to the nearly elliptical shape of E. gigantea. E. pan is most similar
in sculpture, form, and color to E. aurata (Morelet). Differences be-
tween the two species are discussed under the latter.
Euglandina pan has been recorded erroneously in the literature as E.
sowerbyana. Martens (1890:55) recorded a specimen from Guatemala
City, and Jousseaume (1878:169) reported the species from Verapaz. These
two records probably refer to E. pan.
ETYMOLOGY.-The species name pan is from the Classical Greek my-
thology, pan, a deity of the woodland.

Euglandina aurata (Morelet)

Figures 30-31
Glandina aurata Morelet 1849:12.-Fischer and Crosse 1870:106; pl. 3, figs. 7, 7a.-
Martens 1890:57.
Oleacina aurata (Morelet) Gray 1855:33.-Pfeiffer 1859:642.-Tryon 1885:36; pl. 6, fig.
Achatina lignaria Reeve 1849: pl. 8, fig. 27.-Pfeiffer 1853:517.
Oleacina lignaria (Reeve) Gray 1855:34.--Pfeiffer, 1859:642.
Euglandina aurata (Morelet) Pilsbry 1907:188.
DESCRIPTION.-Medium-sized, attaining a length of up to 94 mm.
Elongate-fusiform in shape; about 0.41-0.44 times as wide as high. Spire
equal to or less than height of aperture, being about 0.39-0.52 times
length of shell. Whorls 6.6-7.5; weakly convex with a shallow suture.
Embryonic whorls 3.7, smooth (Fig. 6). First whorl raised and nearly
conical; following two whorls regularly descending. Postembryonic whorls
sculptured with incremental striations that form moderately small white
denticles that crenulate the suture (Fig. 15). Denticles demarked as a
narrow band by a spire groove at their bases. Incremental striations crossed
by weaker spiral striations that form a fine pattern of rectangular granules
which are about half again as high as wide. Spiral sculpture continuous
but diminishing in intensity near base of shell. Aperture elongate-auri-
culate in shape, about 0.48-0.61 times length of shell. Columella rela-
tively long compared to E. pan, straight and vertical, not twisted to the
right (Fig. 30) or forward (Fig. 31). Outer peristome nearly straight in
lateral profile and recurved near base. Color yellowish with irregularly
spaced darker orange streaks along line of growth. Interior of aperture
Measurements in mm of four specimens follow:

VOL. 30 NO. 2


length width height whorls
UMMZ 136471 74 32 39 7.4
USNM 162302 61 27 37 6.6
MCZ 6752 65 27 35 6.5
AMNH 55140 78 34 38 7.5

TYPE LOCALITY.-"Woods of Vera Paz." Undoubtedly this is within
the Departamento Alta Verapaz, near Coban, and not the small village
of Verapaz in the Departamento Esquintla along the Pacific coast. Mo-
relet visited Coban, and this area is within the known range of the spe-
cies. The holotype is in the Museum National d'Histoire Natural in Paris.
DISTRIBUTION (FIG. 32).-This species is known only from higher
elevations in the Departamentos Alta Verapaz and Chimaltenango, Gua-

SPECIMENS EXAMINED.-GUATEMALA-Depto. Alta Verapaz: no fur-
ther data (UMMZ 136471). Depto. Chimaltenango: Santa Elena, near
Tecpan (FMNH 13627). Specimens with no data: AMNH 55410(1), USNM
162302(1), MCZ 6752(1). Martens (1890:57) recorded the species from
Coban and from mountain forests between Tepan (Tecpan?) and Totoni-
capan, 8000-9000 ft alt.
REMARKS.-Euglandina aurata is similar in size and general appear-
ance to E. pan. E. aurata differs from the latter by its finer spiral sculp-
ture which produces finer and shorter granules with the incremental
striations. The two species differ conspicuously in the crenulate denticles
along the suture. In E. aurata the denticles are nearly uniform in size
and spacing, they are set off from below by a spiral incised line, and the
suture follows a regular course in its descent. In E. pan the denticles are
irregular in size and spacing, they are not demarcated as a band by a
spiral groove, and the descent of the suture follows an irregular course.
The columella margins of the aperture of the two species differ. E. aurata
has a relatively long, straight columella that is aligned with the axis of
the shell. &. L has a short twisted columella that is flexed to the right
and forward. Both species differ from the other discussed in this paper
by their relatively small size, details of the sculpture as are set forth in
the descriptions, and aspects of the aperture shape and columella.

The names Glandina aurata Morelet 1849 and Achatina lignaria Reeve
1849 are based on nearly identical specimens. Both names were pro-
posed at the same time but aurata has page priority over lignaria.


Euglandina titan, new species

Figures 23-24
DESCRIPTION.--Shell gigantic, reaching a length of over 110 mm.
Obese, elongate ovate in shape, about 0.50 times as wide as high, with a
corpulent body whorl. Spire elongate, almost as high as aperture; first
five whorls of spire forming a concave outline; subsequent part of spire
weakly convex. Whorls 8.0; suture moderately impressed; whorls evenly
convex between sutures. Embryonic whorls 2.2, smooth (Fig. 4). First
whorl protruding and conical; following whorls regularly descending. First
two postembryonic whorls with low, evenly spaced riblets that terminate
abruptly at suture. Sculpture on subsequent whorls smooth and glossy.
Suture weakly crenulate by a subsutural cord of enlarged irregular poorly
defined beads (Fig. 14). Surface of whorls with weak growth striations
and occasional minute papillae that may be aligned in short spiral or
longitudinal series (Fig. 14). Spiral striations absent. Aperture narrowly
auriculate; columella weakly spiral, truncate, slightly curved to the right.
Outer lip slightly sinuous in lateral profile. (The periostracum of the hol-
otype is completely denuded from the exterior surface; a portion of the
parietal callus 2 cm long was removed from deep within the aperture to
expose the underlying periostracum.) Color light orange with occasional
longitudinal reddish streaks such as occur in E. sowerbyana. Measure-
ments of the unique holotype are: length, 112 mm; width 57 mm; aper-
ture height 62 mm. Type Locality.-Guatemala, Departamento Izabel,
Montanas del Mico, 4 km WSW of Puerto Santo Tomas at 800 m eleva-
tion in a tropical rain forest. Holotype: UF 35307; collected 2 May 1981
by Jonathan A. Campbell.
DISTRIBUTION (FIG. 32).--Known only from the type locality in the
Montanas del Mico, Guatemala.
REMARKS.-This species is similar in shape and size to Euglandina
sowerbyana (Pfeiffer) and E. gigantea Pilsbry. The three species are huge,
exceeding 100 mm in length. The general shapes of the shell are similar.
They also have similar coloration, although E. titan is darker. E. titan
differs from the other two species by its more ponderous shape, larger
number of whorls, smooth sculpture, and nearly straight columella. Both
E. sowerbyana and E. gigantea are narrower, being about 0.40-0.48
times as wide as high, both have a maximum of 7.5 whorls, both have
decussate sculpture consisting of coarse longitudinal and spiral striations,
both have a distinct, beaded subsutural cord that crenulates the suture,
and both have a spiral columella that is strongly flexed to the right. The
sparce, weakly papillate sculpture of E. titan appears to be an extreme
modification of the decussate sculpture of the other two species.

VOL. 30 NO. 2


ETYMOLOGY. -Titan from the classical Greek mythological diety Ti-
tan, the neuter child of Uranus and Gaea, a symbol of large size and
fearful nature, alluding to the snail's gigantic size. It is the largest known
terrestrial carnivorous snail.


Albers, J. C. 1850. Die Heliceen, nach naturlicher Verwandtschaft systematish geordnet.
1-262. Berlin.
Baker, H. B. 1941. Outline of American Oleacininae and new species from Mexico. Nau-
tilus 55:51-61; pl. 5.
1943. The mainland genera of American Oleacininae. Proc.Acad. Nat. Sci. Phil-
adelphia 95:1-14; pls. 1-3.
Beck, H. 1837-1838. Index molluscorum prasenti aevi musei principis augustissimi Chris-
tiani Frederici. 1-124. Hafniae.
Crosse, H., and P. Fischer. 1869. Diagnoses Molluscorum novorum Guatemalae et Re-
publicae Mexicanae. J. Conchyl. 17:28-36, 190-192, 250, 251, 423-426.
Fischer, P., and H. Crosse. 1870-1878. Etudes sur les mollusques terrestres et fluviatiles
du Mexico et du Guatemala. Vol. 1:1-702. Paris.
Gray, J. E., and L. Pfeiffer. 1855. Catalogue of the the pulmonate or air-breathing Mol-
lusca in the collection of the British Museum, 1:1-173. London.
Jousseaume, F. 1878. Excursion malacologique atraves l'exposition de 1878. Bull. Soc.
Zool. France 3:164-178.
Lea, I. 1834. Observations on the naides, and descriptions of new species of that and
other families. Trans. Amer. Phil. Soc. V:23-85.
Martens, E. von. 1863. Uber neue Mexikanische Landschnecken. Monatsberichte der
Koniglich Preussischen Akademie der Wissenschaften zu Berlin (1883). 1884:540-542.
- 1890-1901. Biologia Centrali-American. Land and Freshwater Mollusca. i-xxviii,
1-706; pls. 1-44. London.
Morlet, A. 1849. Testacea novissima Insulae Cubae et Americae Centralis. 1-92. Paris.
Pfeiffer, L. 1846. Dianosen neuer Heliceen. Zeitschrift fur Malacozoologie. 3:158-160.
- 1853. Monograph heliceorum viventium. 111:1-711. Leipzig.
-- 1859. Monograph heliceorum viventium. IV:1-920. Leipzig.
Pilsbry, H. A. 1907-1908. Manual of Conchology, Ser. II, vol. 19:1-366; pls. 1-52. Phila-
1926. Costa Rican land shells collected by A. A. Olsson. Proc. Acad. Nat. Sci.,
Philadelphia, 78:127-131; pl. 10, figs. 8-9; pl. 11, Figs. 1-7.
Strebel, H. 1875. Beitrag zur Kenntniss der Fauna Mexikanischer Land- und Susswasser-
Conchylien, 11:1-58; pls. 1-13.
- 1878. Beitrag zur Kenntniss der Fauna Mexikanisher Land- und Susswasser-Con-
chylien, 111:1-51; pls. 1-22.
Tryon, G. 1885. Manual of Conchology. Ser. II, Vol. 1:1-364; pls. 1-60. Philadelphia.



Figures 1-6.-Embryonic whorls of Euglandina. (1) E. s. sowerbyana (Pfeiffer) (UF 20807).
(2) E. gigantea Pilsbry (UF 20751). (3) E. vanuxemensis (Lea) (UF 34571). (4) E. titan,
n. sp. (UF 35307). (5) E. pan, n. sp. (USNM 487385). (6) E. aurata (Morelet) (UMMZ

VOL. 30 NO. 2


7 8 9

Figures 7-9.-Euglandina vanuxemensis (Lea). (7) (UF 34909). (8-9) (UF 20808).


10 11

12 13 ;

14 15

Figures 10-15.-Sculpture of Euglandina. (10) E. S. sowerbyana. (11) E. vanuxemensis
(Lea) (UF 34709). (12) E. gigantea Pilsbry (UF 20751). (13) E. pan, n. sp. (UMMZ
467385). (14) E. titan, n. sp. (UF 35307, Holotype). (15) E. aurata (Morelet) (UMMZ

VOL. 30 NO. 2


Figure 16.-Distribution of three species of Euglandina in Mexico. E. vanuxemensis (Lea).
E. s. sowerbyana (Pfeiffer). E. s. estephaneae (Strebel).





20 f

Figures 17-22. Euglandina sowerbyana (Pfeiffer). (17-18) E. s. sowerbyana (Pfr.) (UF
21334). (19-20) E. s. sowerbyana (Pfr.) (USNM 317426). (21) E. s. estephaneae (Stre-
bel) (UMMZ 10969). (22) E. s. estephaneae (Strebel) (FMNH 75).

VOL. 30 NO. 2


23 24

25 26

Figures 23-26.--Euglandina. (23-24) E. titan, n. sp., Holotype (UF 35307). (25) E. gi-
gantea Pilsbry (UF 20751). (26) E. gigantea Pilsbry (UMMZ 28711).


Figure 27.-Distribution of Euglandina gigantea Pilsbry in Costa Rica and Panama.

VOL. 30 NO. 2





29 A


30 F 31

Figures 28-29.-Euglandina pan, n. sp. Holotype (USNM 467385). Figures 30-31.-
Euglandina aurata (Morelet) (UMMZ 136471).


Geographic distribution of
Euglandina based upon
specimens examined
A E. titan E. aurata 0 E. pan

Figure 32.-Distributions of three species of Euglandina in Guatemala.

VOL. 30 NO. 2

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