Content uploaded by Michel Dominguez
Author content
All content in this area was uploaded by Michel Dominguez on Mar 08, 2016
Content may be subject to copyright.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit
publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to
critical research.
Taxonomy of the Blind Snakes Associated with Typhlops
lumbricalis (Linnaeus, 1758) (Scolecophidia, Typhlopidae) from
the Bahamas Islands and Cuba
Author(s): Michel Domínguez and Raúl E. Díaz, Jr.
Source: Herpetologica, 67(2):194-211. 2011.
Published By: The Herpetologists' League
URL: http://www.bioone.org/doi/full/10.1655/HERPETOLOGICA-
D-10-00024.1
BioOne (www.bioone.org) is an electronic aggregator of bioscience research content,
and the online home to over 160 journals and books published by not-for-profit societies,
associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content
indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/
terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial
use. Commercial inquiries or rights and permissions requests should be directed to the
individual publisher as copyright holder.
TAXONOMY OF THE BLIND SNAKES ASSOCIATED WITH TYPHLOPS
LUMBRICALIS (LINNAEUS, 1758) (SCOLECOPHIDIA, TYPHLOPIDAE)
FROM THE BAHAMAS ISLANDS AND CUBA
MICHEL DOMI
´NGUEZ
1,5
AND RAU
´LE. DI
´AZ,JR.
2,3,4
1
Centro Iberoamericano de la, Biodiversidad, (CIBIO), Universidad de Alicante, Edificio de Ciencias III, Campus San
Vicente del Raspeig, 03080, Alicante, Spain
2
University of Kansas Medical Center, Kansas City, KS 66160, USA
3
Stowers Institute for Medical Research, Kansas City, MO 64110, USA
4
University of Kansas Natural History Museum and Biodiversity Research Center, Lawrence, KS 66045, USA
ABSTRACT: The taxonomic status of Typhlops lumbricalis is discussed. Populations of Typhlops lumbricalis
(sensu stricto) are redescribed and restricted to the Bahamas islands. Two new Cuban species associated with
T. lumbricalis are described. The new species are medium in size; rostral in dorsal view is a narrow to broad
oval, preocular contacting third supralabial only, low middorsal scale count (,300), 20 scale rows anteriorly
and reducing to 18 posteriorly at around midbody. These can be placed within the T. lumbricalis species
group and a key to the West Indian species is presented.
RESUMEN: La situacio
´n taxono
´mica del nombre Typhlops lumbricalis es discutida. Las poblaciones de
Typhlops lumbricalis (sensu stricto) son redescritas y restringidas a las Bahamas. Dos nuevas especies
cubanas asociadas con T. lumbricalis son descritas. Las nuevas especies son de mediano taman
˜o, con rostral
en vista dorsal desde estrecha a oval amplia, preocular contactando solamente con la tercera supralabial, bajo
conteo de escamas mediodorsales (,300), 20 hileras de escamas anteriormente, reduciendo a 18
posteriormente alrededor del medio del cuerpo. Estas especies pueden ubicarse dentro del grupo de especies
de T. lumbricalis y una clave para las especies antillanas y Bahamas es presentada.
Key words: Cuba; Scolecophidia; Serpentes; Taxonomy; Typhlops lumbricalis species group; West Indies
THIS PAPER is the second result of an
exhaustive revision to elucidate the taxonomic
status of the Cuban snakes of the genus
Typhlops. In the first paper, a new species,
Typhlops golyathi (T. biminiensis species
group), was described (Domı
´nguez and Mor-
eno, 2009). This addition raised the Typhlops
species diversity from Cuba to nine; eight
from the T. biminiensis species group and one
from the T. lumbricalis species group (Thom-
as and Hedges, 2007; Domı
´nguez and Mor-
eno, 2009, respectively).
Previously, Cuban Typhlops were associat-
ed with the T. lumbricalis species group and,
despite having evident morphological varia-
tion, were assigned to Typhlops lumbricalis
(Linnaeus, 1758). However, during the revi-
sion of Cuban specimens under the name T.
lumbricalis deposited in the zoological collec-
tions of Instituto de Ecologı
´a y Sistema
´tica,
Cuba, the authors found populations with
morphological differences, meriting distinct
species recognition. Thus in this paper we
elucidate the taxonomic status of the popula-
tions previously recognized as T. lumbricalis
and we describe two new species from Cuba.
MATERIALS AND METHODS
The specimens (n553, all museum speci-
mens) were examined under a binocular stereo-
microscope, MBC-10, and measurements were
taken following Thomas (1989), Wallach (1998),
Thomas and Hedges (2007), and Domı
´nguez
and Moreno (2009). The sex could be identified
in some specimens, dissecting the tail in the
ventral midline (checking for presence or
absence of hemipenes). Total middorsal scale
counts were made in dorsal view from rostral to
tail tip. Longitudinal scale rows were counted
anteriorly (at neck level, 10 scales posterior to
the rostral) and posteriorly (10 scales anterior to
the vent). If reductions in the number of
longitudinal scale rows were seen, we located
the exact body regions where these phenomena
occurred. These regions are visible ventrally,
each reduction consisting of two neighboring
transverse scale rows that fuse and continue as
one longitudinal scale row; this event may take
5
CORRESPONDENCE: e-mail, micheldd@hotmail.es
Herpetologica, 67(2), 2011, 194–211
E2011 by The Herpetologists’ League, Inc.
194
place several times. The last fusion was consid-
ered as the body region where the total
reduction occurs. Place of the reduction is the
distance from snout to the exact region where it
happened; it is presented as the percentage of
total length (% TL) and as the cardinal-
numbered middorsal scale coincident with an
imaginary line perpendicular to the body’s
longitudinal axis that touches the scale fusion.
Dorsocaudal and ventrocaudal scale counts are
defined as the number of scales along the tail in
the middorsal and midventral axes, respectively.
Dorsocaudals were counted starting from an
imaginary line perpendicular to the body’s
longitudinal axis at vent level. The dorsocaudal
counts within a sample are generally less variable
than the ventrocaudal counts, which are often
irregularly arranged (Wallach, 2004).
Supralabial imbrication and head scutellation
patterns follow Wallach (1993 and 2003, respec-
tively). Snout–vent length (SVL) and tail length
(TA), measurements were taken with a ruler (to
1 mm); anterior body diameter (ABD, at neck
level immediately posterior to occiput), midbody
diameter (MBD), posterior body diameter
(PBD, immediately before vent) and midtail
diameter (TD) were measured with Vernier
calipers (to 0.05 mm), and the majority of head
measurements were taken with a 38 measure-
ment ocular coupled to the binocular stereomi-
croscope (to 0.025 mm, except for head length:
to 0.05 mm). Other measurements taken were
total length (TL), head length (HL, from tip of
the snout to posterior margin of nuchal scale),
snout–eye distance (SED, from tip of the snout
to interorbital axis through the middle of both
eyes, in dorsal view), rostral–eye distance (RED,
from posterior rostral apex to interorbital axis
through the middle of both eyes, in dorsal view),
head width maximum (HWM, at nuchal scale
level, in dorsal view), midhead width (HWE, at
eye level, in dorsal view), anterior head width
(HWN, at nostril level, in ventral view),
interorbital distance (IOD, distance between
the middle of each eye), internostril distance
(IND, distance between the innermost margins
of each nostril, in ventral view), length of the
dorsal rostral (RLD, the length of the rostral
from tip of the snout to the its posterior margin
in dorsal view), width of the dorsal rostral
(RWD, widest part of the rostral in dorsal view),
anterior postnasal width (APNW, the distance
between the anterior apex of the preoculars, in
dorsal view), posterior postnasal width (PPNW,
the distance between the inner apex of preo-
culars, in dorsal view), ocular minimum length
(RA, the distance from the posterior most part of
the preocular–ocular suture to the level of the
posteriormost margin of the ocular, in lateral
view), ocular maximum length (RB, the distance
from the anteriormost part of the preocular–
ocular suture to the level of the posteriormost
margin of the ocular, in lateral view), ocular
length mean (OLM, a mean between RA and
RB), ocular sinuosity (OS, a ratio of RA to RB),
ocular width (OW, the distance between
superiorandinferiorocularapicesinlateral
view, perpendicular to body axis), preocular
minimum length (RC, the distance from the
anteriormost part of the preocular to the level of
the anterior most part of the preocular–ocular
suture, in lateral view), preocular maximum
length (RD, the distance from the anteriormost
part of the preocular to the level of the posterior
most part of the preocular–ocular suture, in
lateral view), preocular width (PW, the distance
between superior and inferior preocular apices
in lateral view, perpendicular to body axis),
preocular sinuosity (PS, a ratio of RC to RD),
eye diameter (ED, diameter of the left eye in
lateral view, parallel to body axis), and preocu-
lar–third supralabial suture length (PSL, the
length of the suture between the preocular and
third supralabial, in lateral view). According to
Thomas (1989), the postnasals are described as
being divergent if their edges diverge anterior to
posterior, and parallel or calyculate if they are
bowed (parenthesis-like).
Comparisons among other West Indian
species of Typhlops were made by examining
preserved material in Cuba, the United States,
and France (Appendix I) and using published
data about scutellation and head pattern (Leg-
ler, 1960; Schwartz and Henderson, 1991;
Thomas, 1989; Thomas and Hedges, 2007;
Wallach, 1998). Acronyms follow standardized
usage (Leviton et al., 1985) with the addition of
Colecciones Zoolo
´gicas de la Academia de
Ciencias de Cuba (CZACC), deposited at the
Instituto de Ecologı
´a y Sistema
´tica (IES), Cuba,
and ‘‘Charles T. Ramsden’’ (CTR) historical
collection, deposited at the IES. Localities for T.
lumbricalis were obtained from some museum
databases; we used HerpNet (2008) to access
June 2011] HERPETOLOGICA 195
data on museum holdings of T. lumbricalis from
CM, MCZ, and KU. Geographic coordinates
arebasedontheWGS84datum.
We analyzed the data using Statistica
version 7.0 for Windows. For all quantitative
measurements, we obtained the mean, 61
standard error, and range. All data were
normally distributed (tested using Kolmo-
gorov–Smirnov tests) and had homogeneous
variances (tested using Levene tests). Thus
several one-factor ANOVAs (see variable list
in Table 1) were employed to compare among
the two new Cuban species, T. lumbricalis
(sensu stricto) and Typhlops cf. silus. We also
examined the presence of sexual dimorphism
for all variables stated in Table 1 (except for T.
lumbricalis [sensu stricto], because its sex was
not identified).
TAXONOMY
The taxonomic history of Typhlops lumbri-
calis is complex and spans more than 250 yr,
and a remark on this history is necessary. The
nominal species Anguis lumbricalis was estab-
lished by Linnaeus (1758: 228). The character-
ization of this species by Linnaeus (1758, 1766)
in the 10th and 12th editions of the Systema
Naturae is ‘‘230–7. Color albido flavescens,’’
with its provenance from America. The de-
scription was preceded by three earlier refer-
ences: (1) the Thesaurus of Seba (1734: Vol. I,
137, Tabula 86, fig. 2), (2) the History of
Jamaica of Browne (1756: 460, Tabula 44,
fig. 1), and (3) the Museum Ichthyologicum of
Gronovius (1756: Vol. II, 52, no. 3). Later, this
species was included in a new genus, Typhlops,
and it is the type species (Oppel, 1811).
Following this taxonomy, several authors
have contributed to the understanding of
the morphology and geographic distribution
of T. lumbricalis (sensu lato), restricting its
distribution to South America first, and later
to the West Indies (e.g., Barbour and
Ramsden, 1919; Bibron, 1843; Boulenger,
1893; Dume
´ril and Bibron, 1844; Gundlach,
1880) although Cochran (1924) redefined
the name T. lumbricalis. She commented
that Linnaeus’ type material, with low
middorsal scale row counts, is sufficient to
apply this name only to the blind snakes
with 20 scales rows anteriorly and low
middorsal scale row counts, which encom-
passes specimens from the Bahamas, Cuba,
and Hispaniola. Cochran never mentioned
the possibility of having collected more than
one species despite having found high
middorsal scale row counts in Cuban pop-
ulations (255–340 scale counts in Cuban
individuals) as well as geographic variation
in scale counts between populations (255–
285 in eastern Cuba and 270–320 in Havana
and Pinar del Rı
´o provinces, in western
Cuba). Cranial scutellation differences were
also observed among the Cuban populations
(e.g., variability in the size of the parietals).
After the publication of Cochran (1924),
seven new species belonging to the ‘‘T. lumbri-
calis species group’’ were described from
Hispaniola: T. capitulatus Richmond, 1964; T.
gonavensis Richmond, 1964; T. hectus Thomas,
1974; T. schwartzi Thomas, 1989; T. tetrathyr-
eus Thomas, 1989; and T. titanops Thomas,
1989. Thomas (1989) restricted the name T.
lumbricalis to the populations on the Bahamas
and Cuba, also he concluded that those
Typhlops populations having 20 scale rows
anteriorly, reducing to 18 rows posteriorly; low
longitudinal scale counts (approximately 300 or
less); a rostral which in dorsal view looks like a
narrow oval; and a collar of pigmentation across
the throat are attributable to T. lumbricalis.Just
as in Cochran (1924), Thomas documented
geographic variation in Cuban populations
showing that the Bahaman T. lumbricalis had
lower middorsal scale row counts (less than 300)
than Cuban populations; he also documented
that Bahaman populations are smaller in overall
size (180 mm maximum TL, 140–160 mm
mode), had a narrower rostral in dorsal view,
and a less divergent postnasal pattern. He
restricted the type locality of T. lumbricalis to
the Bahamas islands due to the T. lumbricalis
type specimens having low longitudinal scale
counts (Linnaeus, 1758), as found in the Baha-
man populations. Additionally, he considered it
reasonable to restrict the type locality to New
Providence Island, because it had long been the
commercial and political center for the Bahamas
islands and because T. lumbricalis occurs there.
However, ‘‘the type locality of a nominal
species-group taxon is the geographical (and,
where relevant, stratigraphical) place of capture,
collection or observation of the name-bearing
type; if there are syntypes and no lectotype has
196 HERPETOLOGICA [Vol. 67, No. 2
been designated, the type locality encompasses
the localities of all of them’’ (article 76.1 from
International Code of Zoological Nomenclature
[ICZN] 1999). Therefore, the ‘‘type locality’’
proposed by Thomas (1989) is not in line with
the criteria set by the ICZN.
Later, McDiarmid et al. (1999, p. 108)
summarized the name T. lumbricalis, included
two synonyms for this name, Typhlops cubae
Bibron, 1843, and Typhlops silus Legler, 1959,
and also remarked that the specimen of Anguis
lumbricalis from Gronovius’ collection is the
holotype, but this designation is incorrect. All
specimens described by Gronovius, Browne,
and Seba, together with any specimens to which
Linnaeus himself had access, are equally
syntypes (see article 72.4.1 from ICZN, 1999).
However, to select a lectotype from the
Linnaeus, Gronovius, Browne, or Seba collec-
tions could be necessary, because Cuban
populations associated with the name T. lum-
bricalis, having 20 anterior scale rows reducing
to 18 posteriorly, differ in scutellation, pattern,
and other characters from T. lumbricalis
populations of the Bahamas. Unfortunately,
the location of those specimens is currently
unknown. Although a lectotype can be fixed by
means of an illustration or description—and
Linnaeus’ own description and those of the
earlier authors cited by him are all of equal
status and together they constitute the basis on
which the name is made available (see article
74.4, ICZN, 1999)—it is very difficult to do so
because these descriptions are very simple. Also,
when A. lumbricalis was published this taxon
was composite of two names currently used for
two distinct species, Typhlops lumbricalis from
the Cuban archipelago and the Bahamas and
Typhlops jamaicensis (Shaw, 1802) from Ja-
maica. The now-nominal species Anguis jamai-
censis was based on two of the references cited
by Linnaeus and the name jamaicensis is a
replacement name for lumbricalis.
For this reason, we will submit an applica-
tion to the Bulletin of Zoological Nomencla-
ture. In this application, we designate a
neotype for Anguis lumbricalis, and also
propose to the International Commission on
Zoological Nomenclature the designation of a
neotype for Anguis jamaicensis. Designation of
the neotypes will fix the identity of the nominal
species T. lumbricalis and T. jamaicensis and
enable resolution of the taxonomy in the Cuban
populations associated with name T. lumbri-
calis. The proposed neotypes for A. lumbricalis
and A. jamaicensis were collected on Eleuthera
Island (Bahamas) and Jamaica, respectively.
With the analyses mentioned previously, we
can now analyze which Cuban populations with
20 anterior scale rows reducing to 18 posteriorly
couldbeassociatedwithT. lumbricalis (sensu
stricto; Bahaman populations). Although T.
lumbricalis occurs widely in the Cuban archi-
pelago, only three populations (two from eastern
Cuba and one from western Cuba) are similar to
the T. lumbricalis (sensu stricto) because they
maintain a smaller size, have a rostral that is
narrow to broadly oval in shape when viewed
dorsally, and low middorsal scale counts (,300).
However, the three Cuban populations differ in
scutellation, pattern, and other characters as
well as with the T. lumbricalis from the
Bahamas. Therefore, the name T. lumbricalis
(sensu stricto) is resummarized and this species
is redescribed and restricted to the Bahamas
islands. Also two Cuban populations similar to it
are recognized as new species and described in
this paper. The third population could be
associated with name Typhlops silus (see below)
and the remaining Cuban populations confused
with T. lumbricalis and having .300 middorsal
scale counts could be new species as well (M.
Domı
´nguez and A. Fong, personal observation).
SPECIES DESCRIPTIONS
Typhlops lumbricalis (Linnaeus, 1758)
(Figs. 1A and 2A; Table 1)
Anguis lumbricalis Linnaeus, 1758: 228;
Laurenti, 1768: 73 (in part, description in-
cludes species from Africa and America).
Typhlopes lumbricalis Schneider, 1801: 339
(unavailable vernacular name).
Typhlops lumbricalis Oppel, 1811: 380;
Merrem, 1820: 158; Dume
´ril and Bibron,
1844: 287; Jan and Sordelli, 1864: Plates IV,
V, fig. 4 (in part; descriptions include several
species from West Indies and Central America).
nec Argyrophis lumbricalis Gray, 1845: 137.
Typhlops lumbricalis Boulenger, 1893: 31
(in part, description includes several species
from West Indies and Guyana); Barbour,
1901: 59 (only Bahaman populations); Bar-
bour, 1914: 322 (in part, description includes
June 2011] HERPETOLOGICA 197
TABLE 1.—Comparison among two new Cuban Typhlops species of the Typhlops lumbricalis species group, Typhlops cf. silus,andT. lumbricalis using selected characters.
Data are presented as mean 61 SE (range). Significant one-way ANOVAs (F) are in bold fonts for emphasis. Means with different letters (bolded for emphasis) differed
significantly (P,0.05, Bonferroni post-hoc test).
Character
Species of Typhlops One-way ANOVAs
lumbricalis,n56oxyrhinus,n58pachyrhinus,n52 cf. silus,n523 F
3,35
P
Dorsal snout pattern Rounded Ogival Rounded Rounded –
Lateral snout pattern Rounded Pointed Rounded Rounded –
Rostral pattern in dorsal view
Narrow oval Yes Yes No No –
Broad oval No No Yes No –
Broad to narrow oval No No No Yes –
Parietals extending towards
the head midline No No Yes Yes
–
Postnasal pattern
Divergent Weakly Strongly Strongly No –
Calyculate No No No Yes –
SVL (mm) 135 67 (119–162) 205 612 (158–244)*208 64 (204–212) 140 64 (107–168)** –
TA (mm) 3 60.2 (3–4) 9 61 (5–13)*760.0 (7) 3 60.3 (2–7)** –
TL(mm) 139 67 (123–166) 214 613 (163–257)*215 64 (211–219) 144 64 (110–175)** –
ABD (mm) 3.9 60.2 (3.3–4.7) 4.4 60.3 (3.1–5.2)*4.5 60.1 (4.4–4.6) 3.4 60.1 (2.5–4.2)** –
MBD (mm) 4.7 60.3 (3.9–5.6) 6.7 60.8 (3.5–9.1)*7.1 60.6 (6.4–7.7) 4.4 60.2 (3.2–5.9)** –
PBD (mm) 4.9 60.3 (4.0–5.9) 5.0 60.3 (3.7–6.0)*5.2 60.2 (5.0–5.3) 3.9 60.2 (2.9–5.7)** –
TD (mm) 2.6 60.1 (2.1–3.0) 3.6 60.3 (2.0–4.7)*4.0 60.2 (3.8–4.2) 2.3 60.1 (1.5–3.8)** –
TL/TA 40.1 62.2
a
(30.8–47.0) 26.7 62.3
b
(19.8–37.5) 30.7 60.6
ab
(30.1–31.3) 46.6 63.2
a
(25.0–89.8) 9.266 ,0.0005
TL/ABD 36.1 61.2
a
(31.9–39.5) 48.4 61.4
c
(44.1–55.3) 47.8 60.2
bc
(47.6–48.0) 41.3 61.0
ab
(33.5–54.7) 10.537 ,0.0001
TL/MBD 29.7 60.8 (28.0–32.6) 34.5 62.4 (26.8–46.6) 30.7 62.3 (28.4–33.0) 33.3 60.9 (26.9–48.6) 1.415 0.255
TA/TD 1.4 60.1
a
(1.0–1.6) 2.4 60.2
b
(1.7–3.2) 1.8 60.1
ab
(1.7–1.8) 1.5 60.1
a
(0.9–2.3) 14.414 ,0.0001
Scale rows anterior 20 20 20 20, 18 –
Scale rows posterior 18 18 18 18 –
Scale reduction
{
%TL <50 54 62 (43–59) 48 63 (46–51) 55 63 (47–73) –
At middorsal scale level – 154 65 (132–168) 128 610 (118–137) 147 65 (123–169) –
Middorsal scale counts 262 63
a
(256–271) 282 64
b
(265–297) 250 67
a
(243–257) 260 62
a
(247–279) 15.065 ,0.0001
Dorsocaudals
{
–1261 (10–14) 13 61 (12–14) 11 61 (9–14) –
Ventrocaudals 12 60.2 (12–13) 13 61 (10–16) 12.5 61.5 (11–14) 11 61 (8–16) –
HWM/HL 1.08 60.03 (1.01–1.21) 1.09 60.04 (0.94–1.28) 1.01 60.07 (0.94–1.09) 1.05 60.02 (0.75–1.23)
{{
0.623
1
0.605
HWE/HWM 0.82 60.01
ab
(0.76–0.84) 0.77 60.02
a
(0.71–0.86) 0.81 60.03
ab
(0.78–0.83) 0.82 60.01
b
(0.76–0.98)
{{
3.095
1
,0.05
HWN/HL 0.53 60.03
b
(0.46–0.64) 0.39 60.03
a
(0.24–0.47)
{
0.41 60.00
ab
(0.40–0.41) 0.46 60.01
ab
(0.35–0.57)
{{
4.691
11
,0.01
HWN/HWM 0.49 60.01
a
(0.44–0.53) 0.35 60.03
b
(0.21–0.44)
{
0.40 60.02
ab
(0.38–0.43) 0.44 60.02
a
(0.28–0.76)
{{
4.349
11
,0.05
198 HERPETOLOGICA [Vol. 67, No. 2
Character
Species of Typhlops One-way ANOVAs
lumbricalis,n56oxyrhinus,n58pachyrhinus,n52 cf. silus,n523 F
3,35
P
IND/HWM 0.37 60.01 (0.33–0.42) 0.35 60.01 (0.31–0.39)
{
0.44 60.02 (0.42–0.47) 0.39 60.01 (0.33–0.59)
{{
2.168
11
0.110
RWD/RLD 0.56 60.02
a
(0.51–0.61) 0.56 60.01
a
(0.52–0.60) 0.84 60.09
b
(0.75–0.93) 0.69 60.02
b
(0.54–0.87) 12.920 ,0.0001
RED (mm) 0.32 60.03 (0.26–0.45) 0.16 60.03 (0.05–0.33) 0.85 60.03 (0.83–0.88) 0.34 60.04 (0.03–0.67) –
PPNW/APNW 0.80 60.02 (0.77–0.89) 0.78 60.02 (0.69–0.88) 0.69 60.04 (0.65–0.73) 0.80 60.02 (0.63–0.91) 1.413 0.255
RC/PW 0.75 60.03 (0.61–0.84) 0.66 60.03 (0.53–0.76) 0.56 60.05 (0.52–0.61) 0.69 60.02 (0.51–0.83)
{{
2.521
1
0.074
PSL (mm) 0.52 60.03 (0.44–0.60) 0.76 60.11 (0.50–1.50) 0.90 60.03 (0.88–0.93) 0.57 60.02 (0.38–0.72)
{{
–
PS 0.86 60.02
b
(0.79–0.90) 0.82 60.03
ab
(0.69–0.90) 0.86 60.01
ab
(0.85–0.86) 0.77 60.01
a
(0.64–0.88)
{{
4.066
1
,0.05
OS 0.83 60.02 (0.75–0.89) 0.84 60.02 (0.71–0.90) 0.86 60.08 (0.78–0.94) 0.76 60.02 (0.59–0.91)
{{
2.734
1
0.059
OLM/OW 0.60 60.01 (0.55–0.65) 0.63 60.02 (0.56–0.75) 0.58 60.03 (0.56–0.61) 0.63 60.01 (0.52–0.75)
{{
0.712
1
0.551
ED (mm) 0.26 60.01 (0.21–0.30) 0.31 60.02 (0.25–0.36) 0.45 60.05 (0.40–0.50) 0.29 60.01 (0.23–0.38)
{{
–
IOD (mm) 2.20 60.09 (1.92–2.47) 0.66 60.03 (0.53–0.76)
{
3.01 60.11 (2.90–3.13) 2.23 60.06 (1.78–2.74) –
SED/HL 0.50 60.01
b
(0.49–0.52) 0.42 60.02
a
(0.31–0.47) 0.46 60.01
ab
(0.45–0.48) 0.46 60.01
a
(0.40–0.53) 6.054 ,0.005
Abbreviations: SVL, snout–vent length; TA, tail length; TL, total length; ABD, anterior body diameter; MBD, midbody diameter; PBD, posterior body diameter; TD, midtail diameter; HWM, head width maximum; HL, head
length; HWN, anterior head width; IND, internostril distance; RWD, width of the dorsal rostral; RLD, length of the dorsal rostral; RED, rostral–eye distance; PPNW, posterior postnasal width; APNW, anterior postnasal width; RC,
preocular minimum length; PW, preocular width; PSL, preocular–third supralabial suture length; PS, preocular sinuosity; OS, ocular sinuosity; OLM, ocular length mean; OW, ocular width; ED, eye diameter; IOD, interorbital
distance; SED, snout–eye distance.
*n57, data for adult specimens only.
** n520, data for adult specimens only.
{Data for T. lumbricalis and 13 specimens of T. cf. silus (KU 47456–68) not available.
{n57, data for CTR 1129 not available.
{{ n522, data for CTR 2347 not available.
1F
3,34
.
11 F
3,33
.
TABLE 1.—Continued.
June 2011] HERPETOLOGICA 199
several species from West Indies and
Guyana); Cochran, 1924: 174–177 (in part,
description includes specimens from Cuba
and Hispaniola); Peters and Orejas–Miranda,
1970: 317 (in part, description includes several
species from West Indies); Thomas, 1976: 28–
31, tables 1–19, figs. 14–17, 64, 75 (in part,
description includes specimens from Cuba);
Thomas, 1989: 409–432; Schwartz and Hen-
derson, 1991: 651; Wallach, 1998: 400;
McDiarmid et al. 1999: 108 (in part, descrip-
tions include specimens from Cuba).
Diagnosis.—Typhlops lumbricalis (sensu
stricto) is the smallest West Indian blind snake
(166 mm maximum TL). This species, three
described species from Hispaniola (T. schwartzi,
T. tetrathyreus, and T. titanops), and two Cuban
species described in this paper are related and
FIG. 1.—Dorsal view of the head scutellation (bar 52 mm) in (A) Typhlops lumbricalis (KU 273756); (B) Typhlops
oxyrhinus (holotype); (C) Typhlops pachyrhinus (holotype); (D) Typhlops cf. silus (CTR 2349).
200 HERPETOLOGICA [Vol. 67, No. 2
define a species group: the T. lumbricalis (see
Discussion). This group can be distinguished
from other West Indian species groups by its 20
scale rows anteriorly, reducing posteriorly to 18
scale rows, low middorsal scale counts (,350,
occasionally ,300), single preocular contacting
with third supralabial only, and two postoculars.
Typhlops lumbricalis (sensu stricto) differs from
some of these species (including new Cuban
species) by its rounded snout (in dorsal and
lateral views), narrow oval rostral in dorsal view
(0.51–0.61 RWD/RLD), weakly divergent post-
nasal pattern, small parietals, and low middorsal
scale counts (,275). The closest species from
Hispaniola is T. tetrathyreus. Both species have
a rounded snout, low middorsal scale counts
(,285), and narrow oval rostral in dorsal view.
But T. lumbricalis (sensu stricto) can be
FIG. 2.—Lateral view of the head scutellation (bar 52 mm) in (A) Typhlops lumbricalis (KU 273756); (B) Typhlops
oxyrhinus (holotype); (C) Typhlops pachyrhinus (holotype); (D) Typhlops cf. silus (CTR 2349).
June 2011] HERPETOLOGICA 201
distinguished from T. tetrathyreus by its dorsal
snout pattern (rounded vs. weakly ogival) and
postnasal pattern (parallel to weakly divergent
vs. calyculate). Because the new Cuban species
were previously confused with T. lumbricalis,
our discussion below will focus mostly on
diagnostic traits among them.
Description.—Adults are small (166.0 mm
maximum TL) with a slightly robust body.
Variation in morphometric variables is sum-
marized in Table 1.
Specimens with head nondistinct from neck
and rounded in dorsal view, dorsoventrally not
depressed, and almost as long as broad.
Rounded snout in dorsal and lateral views.
Rostral in dorsal view like a narrow oval,
slightly broader than long, with a slightly
sharply pointed apex, not parallel-sided and
almost reaching interocular level; ventral
rostral is broad, obliquely sided, with anterior
and posterior constrictions, posterior constric-
tion near the mouth. Nasal divided (nasal
suture complete) with shallow concavity in
posterior margin; broad nostrils, occasionally
visible in dorsal view and closer to preocular
border than rostral border; the short superior
nasal suture and nostrils directed perpendic-
ularly oriented to the head middorsal line and
obliquely oriented (60uangle approximately)
to the body longitudinal axis in lateral view;
inferior nasal suture with a pronounced curve
and contacting in the posterior third of second
supralabial (or in the midline in some
specimens); nasal–preocular suture coincident
with anterior third of third supralabial and
preocular–ocular suture contacting in the
posterior third of third supralabial; postnasal
pattern weakly divergent. Single subtriangular
preocular, higher than broad, with a small
notch in posterodorsal margin and sinuous
preocular–ocular suture, also contacting with
third supralabial only. Polygonal ocular, al-
most as broad as high; black eye, fitting into
preocular superior midpoint and slightly
distant of the snout; two small postoculars
between parietal and fourth supralabial,
superior postocular slightly larger than inferi-
or postocular and with greater contact with
ocular than inferior supraocular. Both posto-
culars have cycloid pattern. Frontal smallest,
but slightly smaller than postfrontal and
supraocular; supraocular obliquely oriented
(60uangle approximately) to the head midline;
two parietals, twice the size of the frontal and
slightly larger than interparietal, interoccipi-
tal, and nuchal; posterior border of the scales
is rounded except in supraoculars and parie-
tals (slightly sharply pointed). Four suprala-
bials with T-III imbrications pattern; second
supralabial twice the size of first, third
supralabial four times the first, overlapping
with preocular and lower edge of ocular,
fourth supralabial largest, 5.5 times the first,
with very rounded posterior margin. Four
infralabials; first almost as broad as second;
third 1.5 times the second; fourth infralabial
largest, three times the first.
Low middorsal scale counts, ,275; 20 scale
rows anteriorly, reducing to 18 posteriorly at
around midbody.
Dorsum and venter colorations are dark
brown and lighter brown (cream), respective-
ly, in alcohol. Snout and head are pigmented.
Tail is pigmented dorsally.
Distribution.—With this new taxonomic
revision, Typhlops lumbricalis is restricted to
the islands of the Bahamas (Fig. 3A). It occurs
in Little Bahama Bank (Water Cay, Little and
Great Ragged islands) and Great Bahama
Bank: Abaco Islands (Great and Little Abaco
islands), Andros Island, Berry Islands (Great
Harbour Cay), Bimini Islands (South Bimini
Island), Cat Island, Eleuthera Island, Exuma
Cays (Great and Little Exuma islands, Pipe
Cay, Staniel Cay), Long Island, and New
Providence Island.
Records of this species from Guyana and
Florida, USA, are based in specimens stored in
museums(e.g., Myers, 1958; Peters and Orejas–
Miranda, 1970). These records are mistakes
because they were based on misidentifications
and erroneous locality data (Dixon and Hen-
dricks, 1979; McDiarmid et al., 1999).
Natural history.—Fossorial lifestyle in soil,
from coastal or near-coastal habitats through
more mesic habitats. This species was found in
shady coppice, xeric scrub, and wooded habitats
as well as in open, cleared areas under rocks in
red soil (Schwartz and Henderson, 1991).
Remarks.—The collar of pigmentation across
the throat, a characteristic unique to T.
lumbricalis (Thomas, 1989), was not observed,
most likely due to having been washed out from
the ethanol preservation.
202 HERPETOLOGICA [Vol. 67, No. 2
FIG. 3.—Localities for Typhlops lumbricalis species group in the Bahamas and Cuba: solid circles, Typhlops
lumbricalis; solid squares, Typhlops oxyrhinus; open squares, Typhlops cf. silus; semisolid squares, T. oxyrhinus and
Typhlops cf. silus; solid triangle, Typhlops pachyrhinus. (A) Bahamas islands and Cuban archipelago; (B) eastern
provinces from Cuba.
June 2011] HERPETOLOGICA 203
Typhlops oxyrhinus sp. nov.
(Figs. 1B and 2B; Tables 1 and 2)
Holotype.—Adult female CTR 2230 from
Bayate (75u219470N, 20u209070W, 338 m ele-
vation), El Salvador Municipality, Guanta
´-
namo Province, Cuba, collected by Lico
Lobaina. Unknown date, although probably
in the middle of 20th century.
Paratypes.—Juvenile CTR 2211, adult males
CTR 2222–23 from San Carlos (75u219470N,
20u099240W, 214 m elevation), Guanta
´namo
Municipality, collected by Charles T. Ramsden
and Lico Lobaina, respectively, same date as
holotype. Adult female CTR 1129 from Ma-
jimiana (75u229000N, 20u229000W, 294 m ele-
vation), Bayate, collected by Lico Lobaina on
17 June 1921. Adult male CTR 1131 from
Colonia Lajas (75u109260N, 20u049450W, 19 m
elevation), ‘‘Los Can
˜os’’ mill (nowadays named
‘‘El Paraguay’’ factory), Guanta
´namo Munici-
pality, collected by Juan Zapata, same date as
holotype. Adult female CTR 2227 from ‘‘Los
Can
˜os’’ mill (75u089340N, 20u039270W, 2 m
elevation), collected by Charles T. Ramsden in
1930. Adult male CTR 2210 from ‘‘Las Cab-
reras’’ farm (coordinates not known), Guanta
´-
namo Municipality, collected by Charles T.
Ramsden on 19 April 1913. All locations in
Guanta
´namo Province, Cuba.
Diagnosis.—A species of medium size
(257 mm maximum TL) of the Typhlops
lumbricalis species group, can be distin-
guished by the combination of its snout
pattern (ogival and pointed in dorsal and
lateral views, respectively) and having more
middorsal scales (.265, 282 mean) than any
other Cuban species of the group and T.
lumbricalis (Table 1). Typhlops oxyrhinus also
differs from other species in this group by its
narrow oval rostral in dorsal view (0.52–0.60
RWD/RLD), postnasal pattern (strongly di-
vergent), robust tail (1.7–3.2 TA/TD), parietal
pattern, and head size.
Description of holotype.—A medium-sized
Typhlops, 232.0 mm SVL, 12.0 mm TA,
244.0 mm TL, robust body, 5.1 mm ABD,
9.1 mm MBD, 5.5 mm PBD, and 3.8 mm TD.
Specimen with head nondistinct from neck
and triangular in dorsal view, dorsoventrally
depressed, broader than long, 0.95 HWM/
HL, narrowing to the apex, 0.83 HWE/HWM,
and 0.38 HWN/HWM. Sharply pointed snout,
narrow, 0.37 IND/HWM. Rostral in dorsal
view like a very narrow oval, longer than
broad, 0.57 RWD/RLD, with slightly sharply
pointed apex, not parallel-sided, and almost
reaching interocular level, 0.33 mm RED;
ventral rostral is broad, obliquely sided, with
anterior and posterior constrictions, posterior
constriction near the mouth. Nasal divided
(nasal suture complete) with shallow concavity
in posterior margin; broad nostrils, occasion-
ally visible in dorsal view and closer to
TABLE 2.—Sexual dimorphism in Typhlops oxyrhinus. Numbers represent mean 61 SE (range).
Character
Sex
Female (n53) Male (n54)
SVL (mm) 216.0 615 (187–232) 196.0 618 (158–244)
TA (mm) 10.0 62 (6–12) 9.0 62 (5–13)
TL/TA 24.8 63.7 (20.3–32.2) 25.6 62.6 (19.8–32.6)
ABD (mm) 4.8 60.3 (4.2–5.1) 4.1 60.5 (3.1–5.2)
MBD (mm) 7.9 61.1 (5.7–9.1) 5.8 61.1 (3.5–8.9)
PBD (mm) 5.5 60.3 (5.0–6.0) 4.6 60.4 (3.7–5.6)
TL/ABD 47.1 60.6 (46.0–47.9) 50.4 62.3 (44.4–55.3)
TL/MBD 29.3 62.3 (26.8–33.9) 37.3 63.6 (28.9–46.6)
TL/PBD 41.1 61.7 (38.6–44.4) 44.6 61.8 (40.0–48.5)
Middorsal scale counts 274 65 (265–283) 287.0 65 (277–297)
Distance of the reduction in the
number of anterior longitudinal
scale rows (given as % TL) 47.5 63.3 (40.9–50.8) 53.6 61.4 (49.7–56.5)
Anterior middorsal scale counts 141.3 65.5 (132–151) 160.5 64.8 (147–168)
Dorsocaudal counts 10.7 60.3 (10–11) 13.0 60.4 (12–14)
Ventrocaudal counts 10 60.3 (6–12) 15.0 60.6 (13–16)
Abbreviations: SVL, snout–vent length; TA, tail length; TL, total length; ABD, anterior body diameter; MBD, midbody diameter; PBD, posterior body
diameter.
204 HERPETOLOGICA [Vol. 67, No. 2
preocular border than rostral border; the
short superior nasal suture and nostrils
directed perpendicularly oriented to the head
middorsal line and obliquely oriented (45u
angle approximately) to the body longitudinal
axis in lateral view; inferior nasal suture with a
pronounced curve and contacting the posteri-
or third of second supralabial; nasal–preocular
suture coincident with second–third suprala-
bials suture; preocular–ocular suture coinci-
dent with third–fourth supralabials suture;
postnasal pattern divergent, 0.75 PPNW/
APNW. Single subtriangular preocular, higher
than broad, 0.53 RC/PW, with a small notch in
posterodorsal margin, contacting with third
supralabial only 0.75 mm PSL; sinuous
preocular–ocular suture, 0.88 PS, 0.82 OS.
Polygonal ocular, slightly higher than broad,
0.65 OLM/OW; black eye, 0.33 mm ED,
2.88 mm IOD, fitting into preocular notch,
and slightly distant from the snout, 0.43 SED/
HL; two small postoculars between parietal
and fourth supralabial; superior postocular has
greater size and contact with ocular than
inferior supraocular, also it is transversely
oriented to body axis, whereas inferior su-
praocular has cycloid pattern. Frontal as
broad as postfrontal and slightly smaller than
supraocular; supraocular obliquely oriented
(15uangle approximately) to the head midline
in dorsal view; small parietals and occipitals
and both scales transversely oriented to
longitudinal body axis in dorsal view, not
extending toward the head midline and similar
to costals; interparietal largest, approximately
1.5 times the frontal, postfrontal, interoccipi-
tal, supraocular, nuchal, and somatic. Posteri-
or border of the scale is very rounded in
supraocular; slightly rounded in frontal, post-
frontal, postparietals, occipitals, and postocu-
lars; and slightly truncated in interparietal,
interoccipital, and nuchal. Four supralabials
with T-III imbrication pattern; second supra-
labial twice the size of first; third supralabial
four times the first, overlapping with preocu-
lar and lower edge of ocular; fourth suprala-
bial largest, 5.5 times the first, with convex
posterior margin. Four infralabials; first as
broad as second; third 1.5 times the second;
fourth infralabial largest, three times the first.
Medium middorsal scale counts, 275; 20
scale rows anteriorly, reducing to 18 posteri-
orly at around midbody: 53% TL or 151th
middorsal scale level. Dorsocaudals, 11, and
ventrocaudals, 10. Three scales border open-
ing of vent. Tail with a small apical spine.
Dorsum coloration is light brown in alcohol,
this is poorly differentiated from venter and
fading over a midlateral zone to lighter venter.
The scales rostral, nasals, preocular, ocular,
and supralabials are slightly unpigmented, tail
is pigmented in dorsal part.
Variation.—All specimens closely approxi-
mate the holotype in scale characteristics and
shape. Variation in the inferior nasal suture
from the holotype state is present in some
paratypes; they have the suture contacting the
midpoint of second supralabial. Paratypes have
four (CTR 1131, 2211) or five (CTR 1129)
scales border opening of vent. Some paratypes
have slightly unpigmented rostral scales nasals,
second and third supralabials, and anterior
portions of preoculars. Juvenile (CTR 2211)
has 146.0 mm SVL, 4.0 mm TA (150 mm TL),
3.4 mm ABD, 3.9 mm MBD, 3.2 mm PBD, and
1.8 mm TD. Other morphological differences
are presented in Table 1.
Sexual dimorphism.—Although measure-
ments from females and males do not differ
significantly, on average, males presented
larger middorsal scale counts, distance to the
exact region where reduction occurred in the
number of longitudinal scale rows and longi-
tudinal scale counts, tail length, and dorso-
caudal and ventrocaudal scale counts, whereas
females were more robust (Table 2).
Etymology.—In Greek oxys 5sharply
pointed and rhinus 5snout, in allusion to
head with sharply pointed snout.
Distribution.—Typhlops oxyrhinus is
known from the Guanta
´namo Province, Cuba
(Fig. 3): El Salvador Municipality (Bayate,
type locality), and Guanta
´namo Municipality
(‘‘Las Cabreras’’ farm, San Carlos, and ‘‘Los
Can
˜os’’ mill).
Natural history.—Fossorial lifestyle under
soil, from sea level through approximately
400 m elevation. Typhlops oxyrhinus inhabits
two different geographic regions: the north-
western coastline of the Guanta
´namo Bay and
the mountains of the Nipe-Yateras area. The
climate in the first region is hot semidesert
without a characteristic rainfall distribution
(with 7–8 or 9–10 dry months). The zonal
June 2011] HERPETOLOGICA 205
vegetation type is the semidesert scrub with
tree-shaped or columnar cacti, arid evergreen
shrub-woods, and open scrubs with a very rich
and diverse xerothermophilous flora. In the
flat valleys of rivers are secondary savannas
and sugarcane plantations. Holotype and one
paratype (CTR 2222) were encountered un-
der the soil of a sugarcane field. Climate of the
Nipe-Yateras area is xerochimeric winter dry
tropical (subhumid type) with one to two or
three to four dry months. The zonal vegetation
types are seasonal evergreen forests, dry
submontane and montane rainforests, and
wet montane rainforests (Borhidi, 1996).
Typhlops pachyrhinus sp. nov.
(Figs. 1C, 2C; Table 1)
Holotype.—Adult male CAZACC 4.11933
from ‘‘El Liberal’’ coffee plantation (coordi-
nates not known), Caban
˜as Municipality,
Sierra del Rosario, Pinar del Rı
´o Province,
Cuba, collected by Eladio Elso in June 1968.
Paratype.—Adult female CZACC 4.19934,
same data as holotype.
Diagnosis.—A species of medium size
(219 mm maximum TL) belonging to the T.
lumbricalis species group, with broader rostral
(0.75–0.93 RWD/RLD) than any other Cuban
species of the group and T. lumbricalis
(Table 1). It can also be distinguished from
others in the group by its low middorsal scale
counts (265–297), rounded snout in dorsal and
lateral views, parietals extending toward the
head midline, postnasal pattern (strongly
divergent), and head size (HWN/HL and
HWN/HWM).
Description of holotype.—A medium sized
Typhlops, 204.0 mm SVL, 7.0 mm TA,
211.0 mm TL; robust body, 4.4 mm ABD,
6.4 mm MBD, 5.3 mm PBD, and 4.2 mm TD
(3.8 mm).
Specimen with head nondistinct from neck
and rounded in dorsal view, dorsoventrally not
depressed, as broad as long, 0.94 HWM/HL,
0.83 HWE/HWM, and 0.43 HWN/HWM.
Snout rounded in dorsal and lateral views,
0.47 IND/HW. Rostral in dorsal view like a
very broad oval, almost as broad as long, 0.75
RWD/RLD, with rounded apex, not parallel-
sided, and not reaching interocular level,
0.88 mm RED; ventral rostral is broad, with
anterior and posterior constrictions, posterior
constriction near the mouth. Nasal divided
(nasal suture complete) with shallow concavity
in posterior margin; broad nostrils, occasion-
ally visible in dorsal view and closer to
preocular border than rostral border; the
short superior nasal suture and nostrils
directed perpendicularly to the head middor-
sal line and obliquely oriented (60uangle
approximately) to the body longitudinal axis in
lateral view; inferior nasal suture with a
pronounced curve and contacting in the
anterior third of second supralabial; nasal–
preocular suture coincident with second–third
supralabial suture and; preocular–ocular su-
ture contacting in the posterior third of third
suprlabalabial; postnasal pattern strongly di-
vergent, 0.65 PPNW/APNW. Single subtrian-
gular preocular, almost as broad as high, 0.61
RC/PW, with a very small notch in postero-
dorsal margin, also slightly sinuous preocular–
ocular suture, 0.85 PS and 0.79 OS contacting
with third supralabial only, 0.88 PSL. Polyg-
onal ocular, almost as broad as high, 0.61
OLM/OW; black eye, 0.40 mm ED, 3.13 mm
IOD, slightly fitting into preocular notch, and
slightly distant of the snout, 0.48 SED/HL;
two small postoculars between parietal and
fourth supralabial, superior postocular two
times the inferior postocular, it also has
greater contact with ocular than inferior
supraocular, both postoculars have cycloid
pattern. Frontal slightly smaller than post-
frontal and supraocular; supraocular perpen-
dicularly oriented to the head midline; two left
parietals, left parietal slightly smaller than
right parietal and slightly oriented obliquely
(45uangle approximately) to the head midline,
occipitals transversely oriented to body axis;
two interparietals; interoccipital slightly larger
than nuchal, similar to frontal and smaller
than frontal and supraocular. Posterior border
of the scales are very rounded in interparietal,
interoccipital, and postoculars; slightly round-
ed in frontal and postfrontal; slightly truncat-
ed in supraocular, parietals, and occipitals.
Four supralabials with T-III imbrication
pattern; second supralabial twice the size of
first; third supralabial four times the first,
overlapping with preocular and lower edge of
ocular; fourth supralabial largest, 5.5 times the
first, with slightly truncated posterior margin.
Four infralabials; first almost as broad as
206 HERPETOLOGICA [Vol. 67, No. 2
second; third 1.5 times the second; fourth
infralabial largest, three times the first.
Low middorsal scale counts, 243; 20 scale
rows anteriorly, reducing to 18 posteriorly at
around midbody: 46% TL or 118th middorsal
scale level. Dorsocaudals and ventrocaudals,
14. Five scales border opening of vent. Tail
with a small apical spine.
Dorsum and venter colorations are dark
brown and lighter brown (cream) in alcohol,
respectively. Snout and head are pigmented.
Tail is pigmented in dorsal part.
Variation.—Differences in some head scale
counts from the holotype state is present in
the paratype, it has single left parietal and
interparietal. It also has three scales border
opening of vent. Other differences are shown
in Table 1.
Sexual dimorphism.—No significant con-
clusions can be made, only one female and
one male available.
Etymology.—In Greek pachy 5thick,
broad and rhinus 5snout, in allusion to head
with very rounded snout.
Distribution.—Typhlops pachyrhinus is
known from the type locality only (Fig. 3A).
However, it may occur elsewhere, such as in
the Sierra Rosario or Sierra de los O
´rganos
(Cordillera de Guaniguanico, Pinar del Rı
´o
Province) because many endemic species
from Cordillera de Guaniguanico occur in
both mountainous zones (such as Anolis
barstchi, Anolis mestrei, Anolis vermiculatus,
and Anolis [Chamaeleolis]barbatus).
Natural history.—Specimens were encoun-
tered under the soil of a secondary forest in
the Sierra del Rosario. This region has a
xerochimeric climate with one to two dry
months in winter. The zonal vegetation types
are seasonal evergreen forests and dry sub-
montane and montane rainforests, although
originally seasonal evergreen tropical forests
predominated. Most of the native vegetation
was replaced by coffee plantations in 1750–
1800, but these fields were abandoned result-
ing in degraded secondary forests (Borhidi,
1996).
DISCUSSION
An important subject to be discussed is the
possibility that two new species could have
previously been described under another
name. One of them is Typhlops cubae Bibron,
1843, type locality Cuba. According to the
author, one specimen of T. cubae (total
number of specimens examined by author is
unknown) has 22 scale rows at midbody, a
rounded snout, and a supralabial pattern
described of the following manner: ‘‘La
segunda labial es penta
´gona, aproximando su
lado anterior de la primera labial, el posterior
de la tercera y sus bordes superiores de la
nasorostral y la maxilar. La tercera labial, mas
grande, tambie
´n penta
´gona y un poco incli-
nada ha
´cia atra
´s, se une con la precedente por
su borde anterior, a
´la siguiente por el
posterior y a
´la maxilar y al ocular por el
superior.’’ Although nomenclature of some
head scales is obsolete, it is possible to
determine that ‘‘nasorostral’’ and ‘‘maxilar’’
names are now known as superior nasal and
preocular, respectively. Therefore, this spec-
imen has the preocular contacting second and
third supralabials, a diagnostic characteristic
of the T. biminiensis species group. But
illustrations of the head scutellation and
description of the rostral: ‘‘… rostral en la
parte mediana, larga, estrecha, un poco
inflada ha
´cia atra
´s, donde concluye en un
borde arqueado …’’ describe a diagnostic of
the feature T. lumbricalis species group. Also,
syntypes of T. cubae examined (MNHN 3218,
3218a) have 209 and 193 mm SVL, 6 and
4 mm TA (215 and 197 mm TL), 20 scale rows
anteriorly, reducing to 18 posteriorly around
at midbody and relatively far posteriorly, and
290 and 281 middorsal scale counts, respec-
tively. Both specimens have rounded snout
and head in dorsal view, broad to narrow
rostral in dorsal view, and preocular contact-
ing third supralabial only, characteristics that
associates these specimens to other species
groups, e.g. the T. lumbricalis species group
or the T. hectus species group (Fig. 4).
With this new analysis, three general
problems were found and it is difficult to
elucidate the taxonomic status of Typhlops
cubae. First, we do not know the total number
of specimens examined by Bibron (1843).
Second, no Cuban specimens were examined
by us within T. biminiensis and T. lumbricalis
species groups that have all of the character-
istics stated by Bibron (1843). Apparently the
description of T. cubae is a combination of the
June 2011] HERPETOLOGICA 207
diagnostic feature of both species groups (T.
biminiensis and T. lumbricalis). Third, syn-
types MNHN 3218 and MNHN 1999.8218 (ex
MNHN 3218a) are examples of an unde-
scribed Typhlops from central and western
Cuba, not T. cubae. These problems lead us to
consider that T. cubae was possibly described
with a specimen series from the T. biminiensis
and T. lumbricalis species groups, of which T.
biminiensis specimens have subsequently
been lost. Therefore it is already necessary
to continue considering to the T. cubae as
nomen dubium, as proposed by Thomas
(1968).
The second name proposed from Cuba is
Typhlops silus Legler, 1959, type locality
Banes, Holguı
´n Province (Fig. 3). Although
this name was considered as a junior synonym
for ‘‘T. lumbricalis’’ by Richmond (1961), this
name can be valid because specimens (iden-
tified as Typhlops cf. silus in this paper, see
appendix; Figs. 1D and 2D; Table 1) col-
lected from some localities in eastern Cuba
(including specimens from Bayate and San
Carlos, Guanta
´namo Province), have common
characteristics with holotype of T. silus (see
Legler, 1959: 105–107) and the majority of
those characters differ from T. lumbricalis and
the two new Cuban species. Those specimens
have 175 mm maximum TL, 20 scale rows
anteriorly, generally reducing to 18 posteriorly
at relatively far posteriorly, 247–279 middorsal
scale counts, rostral in dorsal view like a
seminarrow oval (0.58–0.96 RWD/RLD), not
parallel-sided, rounded snout, preocular con-
tacting third supralabial only, calyculate post-
nasal pattern, and coloration dark brown in
alcohol (Figs. 1D and 2D; Table 1). It is not
the objective of this paper to resurrect the
name of T. silus. For this recognition and
redescription a new paper is necessary (Dom-
ı
´nguez, M. and R. Dı
´az, personal observation).
Key to the Typhlops lumbricalis species
group of the Bahamas, Cuba, and Hispa-
niola.—Thomas and Hedges (2007) described
four new species from Hispaniola associated
with Typhlops hectus Thomas, 1974, and
divided the T. lumbricalis species group in
two groups: T. hectus and T. lumbricalis. New
species were included in the T. hectus species
group (T. agoralionis,T. proancylops, and T.
sylleptor), with T. eperopeus added to the T.
lumbricalis species group. However, this
species has more than 300 middorsal scales
and commonly 20 anterior scale rows, without
posterior reduction; common characteristics
with other taxa of the T. hectus species group,
whereas T. lumbricalis, the new Cuban
species described in this paper and three
Hispaniolan species (T. schwartzi,T. tetra-
thyreus, and T. titanops), have ,300 middor-
sal scale counts and 20 anterior scale rows,
reducing to 18 scale rows posteriorly. There-
fore we included to T. eperopeus in the T.
FIG. 4.—Dorsal and lateral views of the head scutellation (bar 52 mm) in one Typhlops cubae syntype (MNHN 3218).
208 HERPETOLOGICA [Vol. 67, No. 2
hectus species group. We also redefined the T.
lumbricalis species group (Bahamas, Cuba,
and Hispaniola) to include six species to date:
T. lumbricalis,T. oxyrhinus,T. pachyrhinus,
T. schwartzi,T. tetrathyreus, and T. titanops.
The key for these species groups are the
following:
1. Preocular in contact with second and third
supralabials
______________
T. biminiensis species group
Preocular in contact with third supralabial only
___
2
2. Two preoculars
_____________
T. pusillus species group
One preocular
___________________________________________________
3
3. 20 or 22 anterior scale rows, unreduced
(generally) or reducing posteriorly to 20
scale rows
_______________________________________
T. hec-
tus, T. jamaicensis, and T. sulcatus species groups
20 anterior scale rows (rarely 18), reducing
posteriorly to 19 or 18 scale rows
______________________
4
4. Reduction below 20 scale rows occur on
the posterior body, .73% TL; or .300
middorsal scale counts
______________________________
undescribed Cuban populations associated
with ‘‘T. lumbricalis’’, and rarely specimens
of T. hectus and T. sulcatus species groups.
Reduction below 20 scale rows occur
around midbody, ,73% TL (rarely 18
anterior scale rows, without posterior re-
duction); and ,300 middorsal scale counts
__________________________
5(T. lumbricalis species group)
5. Ogival snout in dorsal view
_______________________
________________________________
T. oxyrhinus (eastern Cuba)
Rounded snout in dorsal view
___________________________
6
6. Broad rostral in dorsal view (0.75–0.93
RWD/RLD)
________
T. pachyrhinus (western Cuba)
Narrow or seminarrow rostral in dorsal view
_____
7
7. Narrow or seminarrow oval rostral in dorsal view
___
8
Narrow rostral with parallel sides in dorsal view
___
10
8. Divergent postnasal pattern; parietals not
extending toward the head midline
____________
_____________________________________
T. lumbricalis (Bahamas)
Calyculate postnasal pattern; parietals ex-
tending toward the head midline
_______________________
9
9. Three or four occipitals not extending
toward the head midline
___________________________
________________________
Typhlops cf. silus (eastern Cuba)
Calyculate postnasal pattern; parietals ex-
tending toward the head midline
_______________
____________________
T. tetrathyreus (western Hispaniola)
10. Strongly divergent postnasal pattern; pari-
etals strongly extending toward the head
midline
____________
T. schwartzi (eastern Hispaniola)
Parallel to weakly divergent postnasal pat-
tern; parietals weakly extending toward the
head midline
_______
T. titanops (western Hispaniola)
Acknowledgments.—We are grateful to A. Arango
(Parque Zoolo
´gico Nacional de Cuba) for preserving the
material used; A. Fong (BIOECO) for collecting some of
the examined specimens; A. Herna
´ndez and A
´. Daniel
(IES) for assistance on the geographic references; Y.
Alegre, J. Morales (Museo Nacional de Historia Natural de
Cuba), M. Vegas, and M. Ca
´rdenas for offering related
scientific literature; I. Ineich (Muse
´um National d’Histoire
Naturelle, Paris, France) for loan to V. Wallach (Museum
of Comparative Zoology, Harvard University) the Typhlops
cubae syntypes; A. Gentry, C. McCarthy, K. Way (British
Museum of the Natural History), Rafe Brown, and Linda
Trueb (University of Kansas Natural History Museum and
Biodiversity Research Center) for access to Typhlops in the
collection; S. O. Kullander (Swedish Museum of the
Natural History) and M. Andersen (Natural History
Museum of Denmark) for sharing their knowledge about
zoological material of the Gronovius, Linnaeus, and Seba
collections and to Gentry, McCarthy and Kullander for
their constructive comments; L. A. Ruiz (Conselleria de
Turisme, Generalitat Valenciana, Spain) for constructive
suggestions on the manuscript; and V. Wallach for to
measure syntypes of T. cubae and offering related scientific
literature. MD is deeply grateful to L. V. Moreno (IES) for
sharing his knowledge about Cuban amphibians and
reptiles. We are grateful to editors and the anonymous
reviewer for truly helpful remarks.
LITERATURE CITED
BARBOUR, T. 1901. Batrachia and Reptilia from the
Bahamas. Bulletin of the Museum of Comparative
Zoology 46:55–61.
BARBOUR, T. 1914. A contribution to the zoogeography of
the West Indies, with special reference to amphibians
and reptiles. Memoirs of the Museum of Comparative
Zoology 44:209–359, 1 plate.
BARBOUR, T., AND C. T. RAMSDEN. 1919. The herpetology
of Cuba. Memoirs of the Museum of Comparative
Zoology 47:71–213, 15 plates.
BIBRON, G. 1843. Reptiles. Pp. 122–123. In R. de la Sagra
(Ed.), Historia Fı
´sica, Polı
´tica y Natural de la Isla de
Cuba. Volume IV, Edition in Spanish. Librerı
´ade
Arthur Bertrand, Sociedad de Geografı
´a, Paris, France.
BORHIDI, A. 1996. Phytogeography and Vegetation ecology
of Cuba. 2nd ed. Akade
´miai Kiado
´, Budapest, Hungary.
BOULENGER, G. A. 1893. Catalogue of the Snakes in the
British Museum (Natural History). Volume I. Taylor
and Francis, London, UK.
BROWNE, P. 1756. The Civil and Natural History of
Jamaica. Patrick Browne, London, UK.
COCHRAN, D. M. 1924. Typhlops lumbricalis and related
forms. Journal of the Washington Academy of Sciences
14:174–177.
DIXON, J. R., AND F. S. HENDRICKS. 1979. The wormsnakes
(family Typhlopidae) of the neotropics, exclusive of the
Antilles. Zoologische Verhandelingen 173:1–39.
DOMI
´NGUEZ, M., AND L. V. MORENO. 2009. Taxonomy of
the Cuban blind snakes (Scolecophidia, Typhlopidae),
with the description of a new large species. Zootaxa
2028:59–66.
DUME
´RIL, A. M. C., AND G. BIBRON. 1844. Erpe
´tologie
Ge
´ne
´rale ou Histoire Naturelle Comple
´te des Reptiles.
Volume VI, Comprenant l’Histoire Ge
´ne
´rale des
Ophidiens, la Description des Genres et des E
´spe
`ces
de Serpent Non Venimeux, Savoir: La Totalite
´des
Vermiformes ou des Scole
´cophides, et Partie des
Circuriformes ou Aze
´miophides; en Tout Vingt-cinq
June 2011] HERPETOLOGICA 209
Genres et Soixante-cinq E
´spe
`ces. Librarie Encyclope
´-
dique de Roret, Paris, France.
GRAY, J. E. 1845. Catalogue of the Specimens of Lizards in
the Collection of the British Museum. Trustees of the
British Museum/Edward Newman, London, UK.
GRONOVIUS, L. T. 1756. Museum Ichthyologicum.
Volume II, Sistens piscium indigenorum and quorun-
dam exoticorum, qui in Museo Laurentii Theodori
Gronovii adservantur, descriptiones ordine systematico.
Accedunt nonnullorum exoticorum piscium icones aeri
incisae. Theodor Haak, Leiden, The Netherlands.
GUNDLACH, J. C. 1880. Contribucio
´n a la Erpetologı
´a
Cubana. Imprenta de G. Montiel y Ca, La Habana,
Cuba.
HERPNET. 2008, Available at http://www.herpnet.org.
Accessed: April 3, 2008.
[ICZN] INTERNATIONAL COMMISSION ON ZOOLOGICAL NO-
MENCLATURE. 1999, International Code of Zoological
Nomenclature. 4th ed. The International Trust for
Zoological Nomenclature, London, UK.
JAN, G., AND F. SORDELLI. 1864. Iconographie Ge
´ne
´rale
des Ophidiens. Volume I. Georges Jan and Ferdinand
Sordelli, Milan, Italy.
LAURENTI, J. N. 1768. Specimen Medicum, Exhibens
Synopsin Reptilium Emendatam cum Experimentis
circa Venena et Antidota Reptilium Austriacorum. Typis
Joannnis Thomae Nobilis de Trattnern, Vienna, Austria.
LEGLER, J. M. 1959. A new blind snake (Genus Typhlops)
from Cuba. Herpetologica 15:105–112.
LEGLER, J. M. 1960. Remarks concerning Bahaman blind
snakes (Genus Typhlops). Herpetologica 16:71–72.
LEVITON, A. E., R. H. GIBBS,E.HEAL,AND C. E. DAWSON.
1985. Standards in herpetology and ichthyology: Part I.
Standard symbolic codes for institutional resource
collections in herpetology and ichthyology. Copeia
1985:802–832.
LINNAEUS, C. 1758. Systema Nature Regna Tria Naturae,
Secundum Classes, Ordines, Synonymis, Locis.
Volume I. 10th ed. Reformata, L. Salvii, Holmiae,
Stockholm, Sweden.
LINNAEUS, C. 1766. Systema Nature Regna Tria Naturae,
Secundum Classes, Ordines, Synonymis, Locis.
Volume I. 12th ed. Reformata, L. Salvii, Holmiae,
Stockholm, Sweden.
MCDIARMID, R. W., J. A. CAMPBELL,AND T. A. TOURE
´.
1999. Snake Species of the World. Volume I. Herpe-
tologists’ League, Washington, DC, USA.
MERREM, B. 1820. Tentamen Systematis Amphibiorum.
Johann Christian Krieger, Marburg, Germany.
MYERS, C. W. 1958. A possible introduction of the snake
Typhlops in the United States. Copeia 1958:802–832.
OPPEL, M. 1811. Suite du 1er. Me
´moire sur la classifica-
tion des reptiles. Annales du Muse
´um d’Histoire
Naturelle, Paris [1810] 16:376–393.
PETERS, J. A., AND B. OREJAS–MIRANDA. 1970. Catalogue of
the neotropical Squamata. Part I. Snakes. United States
National Museum Bulletin, Smithsonian Institution
297:viii +1–347.
RICHMOND, N. D. 1961. The status of Typhlops silus
Legler. Copeia 1961:221–222.
RICHMOND, N. D. 1964. The blind snakes (Typhlops)of
Haiti with descriptions of three new species. Breviora
202:1–12.
SCHNEIDER, J. G. T. 1801. Historiae Amphibiorum
Naturals et Literariae. Crocodilos, Scincos, Chamae-
sauras, Boas, Pseudoboas, Elapes, Angues, Amphisbae-
nas et Caecilias. Volume 2. Friederici Frommanni,
Imprimebat, Jena, Germany.
SCHWARTZ, A., AND R. W. HENDERSON. 1991. Amphibians
and Reptiles of the West Indies. Descriptions, Distri-
butions, and Natural History. University of Florida
Press, Gainesville, Florida, USA.
SEBA, A. 1734. Locupletissimi Rerum Naturalium The-
sauri Accurate Description, et Iconibus Artificiosissimis
Expression, per Universam Physices Historiam.
Volumen 1. J. Wetstenium, Gul Smith and Janssonio-
Waesbergios, Amsterdam, The Netherlands.
SHAW, G. 1802. General Zoology, or Systematic Natural
History. Volume 3 (Amphibia), Part 2. Kearsley,
London, UK.
THOMAS, R. 1968. The Typhlops biminiensis species group
of Antillean blind snakes. Copeia 1968:713–772.
THOMAS, R. 1974. A new species of Typhlops (Serpentes:
Typhlopidae) from Hispaniola. Proceedings of the
Biological Society of Washington 87:11–18.
THOMAS, R. 1976. Systematics of the Antillean Blind
Snakes of the Genus Typhlops (Serpentes: Typhlopi-
dae). Ph.D. Dissertation, Louisiana State University,
Baton Rouge, Louisiana, USA.
THOMAS, R. 1989. The relationships of Antillean Typhlops
(Serpentes: Typhlopidae) and the description of three
new Hispaniolan species. Pp. 409–
–432. In C. A. Woods
(Ed.), Biogeography of the West Indies: Past, Present and
Future. Sandhill Crane Press, Gainesville, Florida, USA.
THOMAS, R., AND S. B. HEDGES. 2007. Eleven new species
of snakes of the genus Typhlops (Serpentes: Typhlopi-
dae) from Hispaniola and Cuba. Zootaxa 1400:1–26.
WALLACH, V. 1993. The supralabial imbrication pattern in
the Typhlopoidea (Reptilia: Serpentes). Journal of
Herpetology 27:214–218.
WALLACH, V. 1998. The Visceral Anatomy of Blindsnakes
and Wormsnakes and its Systematic Implications
(Serpentes: Anomalepididae, Typhlopidae, Leptotyph-
lopidae). Ph.D. Dissertation, Northeastern University
of Boston, Boston, Massachusetts, USA.
WALLACH, V. 2003. Scolecophidia miscellanea. Hamadry-
ad 27:227–245.
WALLACH, V. 2004. Typhlops lazelli, a new species of
Chinese blindsnake from Hong Kong (Serpentes:
Typhlopidae). Breviora 512:1–21.
.Accepted: 16 January 2011
.Associate Editor: Christopher Raxworthy
APPENDIX I
Specimens Examined
Typhlops lumbricalis.—BAHAMAS: South Bimini
Island (Bimini Islands): between Nixe’s Harbour and
airport (KU 273724). Eleuthera Island: Alicetown (KU
273740), E. Rock Sound (KU273748, 27375557).
Typhlops sp.—CUBA: Pinar del Rı
´o Province: Pica
Pica (CZACC 4.11985), Vin
˜ales (CZACC 4.5386). Ha-
vana City Province: Nuevo Vedado (CZACC 4.11958),
210 HERPETOLOGICA [Vol. 67, No. 2
El Laguito (CZACC 4.11981), El Respiro, Santa Fe
´
(CZACC 4.11986). Havana Province: Artemisa (CZACC
4.5387). Matanzas Province: Alameda Central (CZACC
4.11960), Santo Toma
´s (CZACC 4.11913–17). Sancti
Spı
´ritus Province: Caguanes (CZACC 4.11979). Cuba
(without more details; MNHN 3218, 1999.8218, syntypes
of Typhlops cubae).
Typhlops cf. silus.—CUBA: Holguı
´n Province: Banes
(KU 47456–68). Granma Province: ‘‘Los Bu
´caros’’ farm
of the Bingham family, Yao (CTR 2369). Santiago de
Cuba Province: Vista Alegre (CTR 2347, 2369), Santa
Marı
´a del Loreto (CTR 4747). Guanta
´namo Province:
Bayate (CTR 2214, 2229, 2373), Belona (CTR 1133),
Guanta
´namo City (CTR 2265), San Carlos (CTR 2221).
DATE OF PUBLICATION
Herpetologica, Vol. 67, No. 2, was mailed on 6 May 2011
June 2011] HERPETOLOGICA 211
