I

June 1995

Asiatic Herpetological ResearchVol. 6, pp. 132-150

A New Genus for the Ramphotyphlops subocularis Species Group(Serpentes: Typhlopidae), with Description of a New Species
VAN WALLACH
Center for Vertebrate Studies, Department of Biology, Northeastern University, Boston, MA 02115, USA
Abstract.

— A new genus, Acutotyphlops,

isestablished for McDowell's Ramphotvphlops subocularis

species group (minus R. willeyi which is transferred to the R.flaviventer group) based on a parietal bone
projection, head shield fragmentation, and a V-shaped lower jaw.

Two species are revalidated, A. infralabialis

and A. solomonis, and a new species is described from Bougainville Island, A. kunuaensis. Acutotyphlops
kunuaensis, represented by more than 250 specimens, is distributed throughout Bougainville, both in the
coastal lowlands and the interior highlands. Sexual dimorphism is present in certain scutellation andproportional characters. At least five species of typhlopids are now known to inhabit Bougainville Island.

Key words: Ramphotyphlops subocularis group, Acutotyphlops, A. kunuaensis, A. subocularis, A.
infralabialis, A. solomonis, Typhlops adamsi, T. bergi, Papua

Introduction

McCoy (1970), McDowell (1974), andHahn (1980) listed three species of Typhlina

Ramphotyphlops) inhabiting
Bougainville Island in the Solomons:Ramphotyphlops braminus, R.flaviventer(= R. depressus fide Wallach, in prep.), andR. subocularis. In his thorough review of
the typhlopids of New Guinea and the(=Solomon Islands, McDowell (1974) defined
the Ramphotyphlops subocularis species
group as lacking a rectal caecum andexhibiting a wedge-shaped lateral snout
profile. He recognized two species in thegroup, Ramphotyphlops subocularis (Waite,
1897) and R. willeyi (Boulenger, 1900),
considering the latter to be the mostprimitive member, scarcely differing from
R. flaviv enter (Peters, 1864) of theRamphotyphlops flaviventer species group
except in the wedge-shaped snout andabsence of a caecum. Ramphotyphlops
subocularis was described as differing from
R. willeyi in the more specialized
fragmentation of the lateral head shields(multiple preoculars and suboculars vs. asingle preocular and no subocular),increased number of midbody scale rows
(26-36 vs. 20-22), and the acute mandibularsymphysis (V-shaped vs. U-shaped).
Ramphotyphlops willeyi also differs from

R subocularis

New Guinea, Bougainville

frontorostral and paired prefrontals on thedorsum of the head (pers. obs.).

McDowell (1974) placed five nominal
taxa in the synonymy of Ramphotyphlops
subocularis, stating that the number of valid
species remained to be determined but based
upon published data there was no evidence
to suggest that more than one variablewasinvolved.
The taxa
with
R.
were
subocularis
synonymized
Waite,
1918,
Typhlops infralabialis
Typhlops solomonis Parker, 1939,
Typhlops bergi Peters, 1948, Typhlops
keasti Kinghorn, 1948, and Typhlops
adamsi Tanner, 1951 (see McDowell, 1974,
for complete synonymy). An examinationof the types of all of the above taxa, plus the
majority of Ramphotyphlops subocularis
speciesmaterial in museum collections indicates that
the R. subocularis complex consists of atleast four valid species (R. subocularis, R.

infralabialis, R

solomonis, and an
undescribed species that forms the topic ofA description of the cranial
this paper).
.

osteology, internal anatomy, and geographic
variation in the Ramphotyphlops subocularis
group, with emphasis on the new species,
will be the subject of a future paper(Wallach, Wong and Meszoely, in prep).

Methods

the presence of aunicameral right lung (vs. multicameral), a

All traceable museum specimens of the

convoluted and multisegmented liver (vs.
straight and unsegmented), and absence ofRamphotyphlops subocularis group (> 325
specimens) were examined, including the.

in© 1995 by Asiatic Herpetological ResearchDue totypes of all nominal taxa.
misidentifications in collections, somespecimens may have been overlooked.
Specimens were examined with an Olympus
binocular dissecting microscope andmeasurements were made to the nearest 0.5mm, including total length (LOA), tail length(TL), midtail diameter (MTD), anterior(ABD), midbody (MBD) and posterior(PBD) body diameters, and diameter in the
nuchal region (ND).

Total middorsals ortransverse scale rows (TSR) were counted
between the rostral and terminal spine,
dorsocaudals (DC) along the midline in aperpendicular plane to the anterior
ventrolateral edge of the vent to the apicalspine, and subcaudals (SC) between thevent and the spine. Five longitudinal scalerow (LSR) counts were made: postcephalicor anterior (ASR) scale rows were counted
at the level of the

midocular level; head length (HL) is distancefrom tip of snout to midocular level. All
diameter measurements were made in either
dorsal or ventral view.
Statistics

were calculated with theMacintosh Statview program.

Mean values

are presented with their standard errors (SE)and ranges (r); CV represents the coefficientof variation.
Probability values in the
Tables refer to the student's f-test of sample
means. In Tables 4-5 data are presented by
sex and both sexes combined; when astatistically significant difference exists
between the means of the male and female
samples, the probability value is given inplace of the data for the sexes combined.
Tables 6-7 summarize only the combined
sex samples as no statistically significant
sexual difference was found to be present.

20th scale caudad of themental, midbody (MSR) rows at midbody,and precloacal or posterior (PSR) rows werecounted at the level of the 10th scale craniadof the anals; two further values were
calculated by adding the midbody andposterior counts (MPSR) and also theanterior,

Vol. 6, p. 133

Asiatic Herpetological Research

April 1995

midbody and
Relative

posterior counts(AMPSR).
length ratio
tail from
of
the
is
the
(TL/LOA)
lengthtailposterior border of vent to tip of apical spine
divided by overall length, body proportion
ratio (LOA/MBD) is overall length divided

by midbody diameter, and tail proportion
ratio (TL/MTD) is length of tail divided by

the midtail diameter.

Due to inconsistencies in the literature,clarification is given for terminology of thehead shields

in

the

Ramphotyphlops

subocularis group as the fragmentation of
cephalic scutes is a diagnostic character.

As

discussed by Parker (1939) and Kinghorn(1948), the proliferation of head shields inthe R. subocularis group has led toconfusion and uncertainty as to correct

homologies. Kinghorn (1948) proposed themost logical system of nomenclature forthese shields and his system is followed
here with minor changes. Proliferation of
the lateral head shields is the result ofdivision of the ocular and preocular shield of
typical typhlopids.

The Ramphotyphlops

Due to state of preservation and injection

subocularis group exhibits two patterns ofof preservative (either lack thereof or
overinjection), not to mention health of theanimal at time of preservation, both bodyand tail proportion figures only approximate
the condition in life and were thus rounded
to the nearest integer (except in the case oftype specimens); carrying out thecalculations to one decimal point infers aprecision that is unrealistic. Care must be
taken especially in interpreting the tailproportion ratios as injection of tail withpreocular (PR) arrangement: a single large
preocular (R. infralabialis and new species)
or a longitudinally divided shield with alarge superior preocular and a smaller
inferior preocular (R. subocularis and R.
Division of the typicalsolomonis).
shield has produced an
oculartyphlopid
anterior (AO) and posterior ocular (O) and
The suboculars aresuboculars (S).
inor
two horizontal rows
onearrangedpreservative probably leads to a distortion of
the true values but the data are presented in
the hope that all tails have been similarly
biased and therefore of comparative value.

Head width (HW) is diameter of head at

between the oculars and supralabials (L) andtermed superior and inferior, and consideredvertically

in

one

to

three

columnsasanterior, middle and posterior. Postoculars(T) are defined as all scales in contact with

TABLE 1
CharacterMUS.

June 1995

Asiatic Herpetological ResearchVol. 6, p. 134

Variation in the holotypes of the Ramphotyphlops subocularis species groupsubocularis

keastisolomonis

infralabialis

adamsibergi

kunuaensis

June 1995

Asiatic Herpetological Research

Vol.6, p. 135

TABLE 2. Qualitative characters of the Ramphotyphlops subocularis species group
Species

infralabialis

DP

LP

RS

SN

IP

cs

AS

June 1995

Asiatic Herpetological ResearchVol. 6, p. 136

Head shield terminology of the Ramphotyphlops subocularis species group. Head of holotype of T.
subocularis (after Waite, 1897): a) dorsal view, b) lateral view, c) ventral view; head of holotype of T.
ventral view; head of holotype of T.
infralabialis (after Waite, 1918): d) dorsal view, e) lateral view,
solomonis (after Parker, 1939): g) dorsal view, h) lateral view; head of holotype of T. bergi (after Peters,
FIG. 1.

AO = anterior ocular, F - frontal, FL = postfrontal, FR = frontorostral, IP = nasal, P = parietal, PF = prefrontal, PR = preocular, PT = postfrontal, R ==Lsupralabial, Ninterparietal,
1948):i)rostral, S

dorsal view.

= subocular, SN = supranasal, SO = supraocular, T = postocular, stippled eye shield = ocular (T.

subocularis) or posterior ocular (T. infralabialis, T. solomonis)

broad base (aptly described as thorn-like by

Waite) and it points upward as the tail is
flexed dorsally. Whether this is an artifactof preservation (injection with preservative)
or a characteristic of the R. subocularis
group is unknown, but this dorsal flexure of
the tail tip was commonly observed in otherspecimens.

The

nostrilishalf-moon

shaped, oriented at 45° to the vertical, andThe SIP is T-0 (N1/SL1,
PrOc/SL2, ASOc/SL3, PtOc/SL4). The
directed laterally.

first three supralabials are subequal in sizeand length while the fourth supralabialismore than twice as deep and long as any of

the other three. The dorsum is uniformly
dark brown while the venter is goldenyellow. A sharp demarcation separates thetwo colors with only an occasional brownscale appearing in thescale row.

uppermost yellow—

Typhlops keasti. The status of T.
keasti (AMS R 12856) is uncertain. Other
than the distinctive depression of the headwith its laterally pointed snout and strongly
tapered head in dorsal aspect there is nothing
to separate T. keasti from R. subocularis.
Kinghorn (1948) erroneously reported theJune 1995

Asiatic Herpetological Research

Vol.6, p. 137

FIG. 2. Head of holotypc of Typhlops subocularis (AMS R2202): a) dorsal view, b) lateral view; head oflateral view; head of holotype of Typhlops
holotype of Typhlops solomonis (IRSNB 2029): c) dorsal view, d)lateral view; head of holotype of Typhlops adamsi (MVZ
dorsalview,
R4609):
e)(AMS
infralabialis

40753): g) lateral view. Bar = 1 mm.

type of T. keasti to possess 32 midbody
scale rows but it has 34 rows with an
identical formula (40-34-30) to that of thetype of Typhlops subocularis. Also, hewith areported an overall length of 285
thediameterof5but
specimen
midbody
overall with a 10.5
now measures 373
diameter. The apical spine of the tailflexes dorsad. On the left side of the head,

mm

mm
mm

mm
the

SIP

is

MSOc/SL3,supralabials.

PtOc/SL4)

with

four

On the right side the second

PrOc/SL2b/ASOc, MSOc/SL3, PtOc/SL4).
This is clearly an anomalous condition: fivesupralabials occur only rarely in the R.
subocularis group and the more primitive TIII SIP is characteristic of the three otherspecies. The dorsum is reddish-brown andis

The eye is dimly visible with a pupil
and the SIP is T-III (N1/SL1, PrOc/SL2,
PrOc/SL3/SOc, PtOc/SL4). The dorsum is
uniformly dark brown (with a grayish
overcast due to premolting condition) while
above.

the venter is gold with a sharp demarcation
between them. Several anals are white andthe terminal spine is directed slightly ventrad

and orange.

T-0 (N1/SL1, PrOc/SL2,

supralabial is divided, resulting in fivesupralabials, and the third supralabial
overlaps the inferior anterior subocular,
forming a T-III SIP (N1/SL1, PrOc/SL2a,the venter

June 1995

Asiatic Herpetological Research

Vol.6, p. 138—

Typhlops infralabialis. The type of T.
infralabialis (AMS R4609; Figs, ld-f, 2e-f)
has a T-III SIP (N2/SL1, PrOc/SL2,
PrOc/SL3/ASOc, PSOc/SL4) and the tailcurves ventrally. The half-moon shaped
nostril is directed laterally and inclined at 45°
to the body axis. The dorsum and venter are
severely faded but a lineate pattern is faintly
visible; the central portions of each scale arebrown with lighter margins. Pigmentation
decreases ventrally so that the lower scalerows are light with a small dark central spot.

gold with occasional scales
A wide nuchal collar

Typhlops adamsi.

pigmented in brown.

(6-10 scales) is confluent with the light colorof the chin and venter. In addition to thetype of Typhlops keasti I have examined
three additional specimens from Papua New

Guinea referrable
11705-06, 11708).

taxon (NMBA
Several specimens ofto thisadamsi (MVZ 40753; Fig. 2g) is similar toTinfralabialis except that the suture
dividing the preocular from the anterior
ocular is incomplete (as in two specimens of
the undescribed species) and the dorsal headprofile is bluntly rounded rather than.

The SIP is T-III (N1/SL1,
AOc/SL3/ASOc,
PrOc/SL2,
PtOc/SL4); thethe undescribed species resemble T. keastiin the acute depression of the head. It seems

pointed.

preferable to consider Typhlops keasti a
synonym of Ramphotyphlops subocularis as

tail tip issuggested by McDowell(1974).

Robb(1966) reported T. keasti to have a
Ramphotyphlops-likt hemipenis but did not
mention the number of coils. If further
material should substantiate other
differences between T. keasti and R.
subocularis, then T. keasti may be regarded
as a separate sympatric species.

Typhlops solomonis.

—The type ofT.

—The type of T.

straight and terminates in a soft

protuberance (possibly an apical spine waspresent but is missing due to damage). The
color pattern consists of a brown dorsum
with lineate effect (only central half of eachscale pigmented), fading to pink ventrally.

Six specimens of the new species from
Bougainville exhibit a condition similar tothat seen in the type of Typhlops adamsi
with the preocular semifused to the anterior
The type of Typhlops adamsi
ocular.
ananomaloustobeappears

solomonis (IRSNB 2029; Figs, lg-h, 2c-d)
is in a premolting condition with a milky
appearance and numerous sloughing scales

Ramphotyphlops infralabialis and is placedin the synonymy of that species.

along various portions of the body, a factor
making an accurate middorsal countTyphlops bergi. The type of T. bergi(UMMZ 95445; Fig. lh) exhibits a T-III
SIP (N1/SL1, PrOc/SL2, PrOc/SL3/ASOc,
PtOc/SL4). The half-moon shaped nostril is
inclined at 45° and directed laterally. The

Prominent gland depressions arepresent along the head shield margins. A

difficult.

large circular nostril is obliquely oriented ina semidivided nasal and just visible from—inferior nasal suture contacts

SL2 near its

June 1995

Asiatic Herpetological ResearchVol. 6, p. 139

TABLE 3. Variation in the ocular shields of the Ramphotyphlops subocularis species group1Vol. 6, p. 140

Asiatic Herpetological Research

TABLE 5. Proportional characters of Acutotyphlops
Species

June 1995

Revalidation ofTaxa

The summarized data in Tables 1-7
demonstrate the distinctness of the four
species here considered valid (R.

subocularis, R. solomonis, R. infralabialis,and the new species). The R. subocularis
group is characterized by six uniquely
derived features unknown among othertyphlopids.

An osteological synapomorphyof all species is the presence of an acuminate
parietal projection separating the posterior
frontals. This median middorsal "spike" ofthe parietal bone, wedged between theposterior portions of the frontals, has notbeen reported in any other scolecophidian.
Examination of six skulls reveals the parietal
spike to extend for the following distancesalong the interfrontal suture: 0.20-0.25 in R.

solomonis, 0.33 in R. subocularis and theundescribed species, and 0.40 in R.
Multiple preocular, ocular
and/or subocular shields (Table 3), a Vshaped lower jaw with 5-7 infralabials
infralabialis.

(Figs,lc,f),

and a frontorostral shieldbordered by a pair of enlarged prefrontals(Figs, la, d, g, i) are other synapomorphiesof the group as here defined with theexclusion of Ramphotyphlops willeyi.
Additional derived characters present in theR. subocularis group (but not exclusivelyso) include lack of a rectal caecum, auropelud-like nuchal expansion such that thegreatest diameter of the body is behind thehead, a high number of longitudinal scalerows (> 26 midbody rows present in 27
species of African and Asian Typhlops and
15 species of Rhinotyphlops), sexual

dimorphism in relative tail length andnumber of subcaudals and dorsocaudals
1985, for Typhlops
Perry,amulticameral right lungvermicularis),(i.e.,(present in Rhinotyphlops and someTyphlops), and a straight unsegmented liver(present also in Rhinotyphlops).

Ramphotyphlops willeyiisthus transferredto the R.flaviv enter species group,

which

possesses a unicameral right lung. It isconsidered a derived member of that groupbased upon the wedge-shaped snout andabsence of a rectal caecum (McDowell,1974).

Vol. 6, p. 141

Asiatic Herpetological Research

June 1995

Due to the uniqueness of the R.
subocularis group, which is distinguishedfrom all other typhlopids by no fewer thansix

synapomorphies

and

has

been

previously suggested as worthy of separate
generic status (Dunn and Tihen, 1944;1974), a new genus isestablished to contain the four species
discussed herein. The removal of these
species, having 26-36 midbody scale rows,
from Ramphotyphlops leaves all membersof that genus but one with 16-24 midbody
scale rows, the sole exception being thePhilippine Rampfiotyphlops cumingii (24-28
rows). The new genus may be known as

McDowell,Acutotyphlops gen. nov.

Type species.

— Acutotyphlops

kunuaensis sp. nov.

Diagnosis.

— Distinguished fromallother typhlopid genera by any of thefollowing characters: a middorsal parietal
spike partially separating the frontal bones, aV-shaped lower jaw, two or more subocularshields, a frontorostral shield bordered by apair of prefrontals, five or more infralabial
shields, and sum of preocular and ocularshields three or more.

—

Etymology. From the Latin acutus,
meaning pointed, in reference to both theparietal spike of the skull and the symphysisof the lower jaw of the four included
species, plus the dorsal and lateral headprofiles of the type species.

—

Acuminate projection of
Description.
bone
separating posterior frontals
parietal
the interfrontal suture; 4-5
0.2-0.4
of
along
teeth
(4 in A. subocularis and A.
maxillarysolomonis; 5 in A infralabialis andundescribed species); 2-4 dorsal foramina in.

nasal bone; cephalic glands confined tobases of shields beneath sutures; widest partof body in nuchal region due to expanded
axial musculature, presumably adaptive forburrowing and similar to condition in the

Uropeltidae fide Gans, 1976, and Gans et
1978; rostral narrow and short,al.,extending halfway to the level of the eyes;
median azygous frontorostral bordered bypaired prefrontals; superior nasal suture

Asiatic Herpetological ResearchVol. 6, p. 142

June 1995

A. subocularis

NEW IRELAND

Q
A

.4.

solomonis.4.

kunuaensis.4.

infralabialis

BOUGAINVILLE

GB IRG1 \ <9

^4^

Gl

FIG. 3.

Distribution of Acutotyphlops n. gen..

\l'\l

<

\N\1

Solid symbols represent specimens examined, open

symbols denote type localities.

incomplete; caudal border of nasal concave;inferior nasal suture contacting second
supralabial; preocular single or divided
longitudinally; ocular single or divided
transversely (if divided, small eye withpresent beneath posteriorsumofocular);preocular and ocular shields
2-7
3-6;
suboculars; 4 supralabials with
fourth 2-3 times as long as deep and twice
as long as any of the other three; 5-7
infralabials; longitudinal scale rows withanterior and posterior reductions; 26-36
midbody scale rows; sum of anterior,midbody and posterior scale rows 88-110;
visible pupil

lower

jaw

V-shaped;

right

lung

multicameral; liver straight and unsegmented(as in Alethinophidia); rectal caecum absent(as in most Alethinophidia); lateral tongue

papillae absent;

Ramphotyphlops-Uke

hemipenis with 3-9 coils in retracted
position; sexual dimorphism in relative taillength and both subcaudal and dorsocaudal
moderate-sizedscalecounts;scolecophidians with maximum length of

400 mm (except A. solomonis at 500 mm)
and moderate length/width ratios of 25-50.

Content.

— Four recognized species:

Acutotyphlops subocularis (Waite, 1897),
including its synonym Typhlops keasti
Kinghorn, 1948; Acutotyphlops infralabialis(Waite, 1918), including its synonyms
Typhlops bergi Peters, 1948, and Typhlops
adamsi Tanner, 1951; Acutotyphlops
solomonis (Parker, 1939); andAcutotyphlops kunuaensis sp. nov.

Distribution.

— Eastern Papua New

Guinea and the Solomon Islands (Fig. 3).
Acutotyphlops subocularis, the northernform, is known from eastern Papua New
Guinea (one record from Morobe Province)
and the Bismarck Archipelago, NSL-1065
m. The two central forms, recorded from
NSL-915 m, include Acutotyphlops
kunuaensis n. sp., endemic to BougainvilleIsland, and A. solomonis, recorded fromeastern Papua New Guinea (one record fromMilne Bay Province) and Bougainville.

Vol. 6, p. 143

Asiatic Herpetological Research

June 1995

Artificial key to the genus Acutotyphlops
la.

Preocular single, anterior suboculars paired, dorsal snout

2

profile roundedlb.

Preocular divided, anterior suboculars single, dorsal snout
3

profile pointed

2a.

Supranasals present, ocular single, supralabial imbrication
pattern T-0

A. subocularis

2b.

Supranasals absent, ocular divided, supralabial imbrication

3apattern T-HI
Midbody scale rows 26-28

A. infralabialis

3b.

Midbody scale rows 30-36

A. kunuaensis

Acutotyphlops infralabialis, the southernform, has the widest distribution, being
known from Bougainville, New Georgia,
Malaita, and Guadalcanal in the Solomon
Islands, 15-245 m.

Specimens of Acutotyphlops have notbeen reported from some of the large islands

Solomons (Choiseul, Florida, San
Thus its
Cristobal, and Santa Isabel).
distribution in the southern Solomons isin thepoorly known. More collecting is urged,
not only in the southern Solomons, but alsoineastern Papua New Guinea as
Acutotyphlops may occur in other localities
along the eastern coast.

Type Species of Acutotyphlops

The species to be designated as the typeof Acutotyphlops has been recognized as anovel taxon for 25 years and is represented
in the

MCZ collection by more than 220

individuals collected by Fred Parker from

August 1960 to May 1966, 180 of themfrom the type locality of Kunua. Several
workers have borrowed the MCZ material to
study but it has never been described. This
blind snake was mentioned by Parker(1970) as being one of 13 new species from
Bougainville that "either have been
described or soon will be," based upon nine
years of collecting by himself and natives.
The new form is finally christened

A. solomonis— MCZ

76964, an adult male
Holotype.
collected by Fred Parker (field no. X-4688)
on 19 August 1963.

Type locality.

— Kunua,

coastalnorthwestern Bougainville Island, North
Solomons Province, extreme eastern Papua
New Guinea, 5°46'S, 154°43'E, elevation
ca. 30 m.

—

Same locality andParatypes (n = 180).
collector as that of the holotype (date ofcollection in parentheses following catalogue

number): MCZ 72067-74 (21.vi.62),
72075-77 (22.vi.62), 72078 (27.vi.62),

72080(22.vii.62),

72130(25.xii.62),

72131-32 (24.V.62), 72133-36 (13.vi.62),
76714, 76716-26 (24.vii.63), 76926-27,
76929-30 (28.vii.63), 76931-32, 76935-39(ll.viii.63), 76950, 76955-57, 76959,
124473 (16.viii.63), 76960 (21.viii.63),
76961-65, 76967 (19.viii.63), 76968-74,
76977, 76979-80, 76982-83, 76986-89
76990-96, 76998-7007(28.viii.63),(29.viii.63), 77008-13 (31.viii.63), 7701622 (5.ix.63), 77023-33, 77036, 77038(8.ix.63), 77037 (15.ix.63), 77267-79,
77282-90 (l.ix.63), and 77292-306(12.ix.63).

Collection date unknown for thefollowing paratypes:

MCZ 76206, 76682-

87, 76690-700, 76704-12, 76948.

—

Etymology. The specific epithet isderived from the type locality, Kunua,
where the entire type series originated.

Acutotyphlops kunuaensis sp. nov.
Figs. 4a-b, 5—

Diagnosis. Distinguished from allother typhlopids by the following
combination of characters: snout pointed in
dorsal and lateral aspects, mandibles V-

Vol. 6, p. 144

Asiatic Herpetological Research

June 1995

FIG. 4. Head of holotype of Acutotyphlops kunuaensis n. sp. (MCZ 76964): a) dorsal view, b) lateral view.
Bar = 1 mm.

Melelupj

Muniha,/*%* \ Topanos^~~~~"vaJune 1995

Asiatic Herpetological Researchanterior subocular plus superior and inferior
posterior suboculars. Bordering posterior
ocular and suboculars, between the
supraocular and SL4, are three small
postoculars on left side and four postoculars
on right. Supralabial imbrication pattern T-

APrOc/SL2,(N1/SL1,

III

APrOc/SL3/PPrOc, PtOc/SL4).

Fourthandtosupralabial elongatedequallength offirst three combined.
All head shields
possess tiny convex tubercles irregularly
scattered over their surfaces.
In ventral

andslightly

view snout acutely pointed
trilobate, and mandibles oflower jaw acutely angled or V-shaped withslightly bulbous symphysis. Six infralabials
present on each side, separated by medianmental shield.
Overall coloration is dark brown dorsum

andventerwithpale yellow
strongdemarcation between the two colors. Only

occasionally are there a few lightly
pigmented scales in uppermost yellow row.
Each dorsal scale pigmented centrally withvery narrow light margin around periphery.
Wide yellow nuchal collar 4-5 scales long
middorsally and 8 scales midlaterally.
Middorsal 18 scale rows (17 posteriorly)
heavily pigmented while yellow rows

Vol. 6, p. 145

between 300 and 380 mm. Females appear
one (n=6) or two (n=5) eggs per
cycle, although one female (MCZ 72136)
has four developing ova on the left side witha large egg on the right. Females with large
eggs were collected on 24 July, 28-29
August and 5 September 1963, indicatingthat egg deposition probably occurs inAugust and/or September.
to lay—Variation.
Lateral tongue papillae areabsent in nine specimens with protruded
tongues (AMS 121957, 123396, 12340203,
76695-96, 76722, 76972,
76986). An inferior nasal pit, similar to thatin the type of Typhlops bergi, is visible in atleast 21 specimens (8.2% of sample).

MCZ

The mean number of helical coils in theretracted hemipenis is 5.9±0. 176 SE in 68
specimens with a range of 3-9 coils andmodal values of 6.5 and 7 (9 specimens
There is a positive but weakeach).
correlation between the number of coils andtotal length,with regression formula for

number of hemipenis coils = 0.02 LOA +1.38 (R 2 = 0.29, CV = 21.2, F = 27.25
with 67 df, p < 0.0001).
The large
variationinthenumberof coils
intraspecificin the retracted hemipenis is surprising andBay, Empress Augusta Bay), east-central(North Nasioi, Kieta) and southern (Buin)urges caution in using the number as asystematic character (McDowell, 1974;
Wallach, 1993b). Variation in hemipenial
coils should be examined in species ofRamphotyphlops as it may be more stable inthat genus. If not, then the number of coils
would appear to have little taxonomic value.
The mean number of coils in the hemipenisof other members of Acutotyphlops is asfollows: A. solomonis - A.l (r = 4.5-5, n =- 5.5 (r = 46, CV = 16.1), A. subocularis

coastal regions plus the northern interior

8, n(Melelup, Mutuhai, and Topanos).

8.2 (roccupy 20-14-1 1 ventral scale rows.
Distribution.

—Endemic

to BougainvilleIsland, Northern Solomons Province, Papua

New Guinea

(Fig.

4).

Apparently

widespread over the island as it is known
from the northwestern (Kunua), westcentral (Cape Torokina, Torokina, Torokina= 8, CV = 24.8), and A. infralabialis = 7.5-9, n = 3, CV = 9.4). In— The population

juvenile specimens the hemipenis is foldedin a zig-zag configuration rather than coiled

structure of Acutotyphlops kunuaensis is

in loops. One specimen (MCZ 77306), witha tail length of 9.5 mm, has both hemipenes

Natural history.

such that juveniles and immature individuals
appear to have total lengths less than 180-220 mm. The smallest gravid females are222
in length. Adults average 220-300
in length. Mean female length is greaterthan that for males. Only 3 males (2.7% of
sample) have total lengths greater than 300
but 16 females (11.3%) have lengths

mmfully everted, the right organ measuring 43
in length and the left one 30 mm. The

mm

hemipenis is nude, 0.5

mm in diameter, andmm

exhibits a sulcus spermaticus extending themmlength of the organ that is V-shaped incross-section with a lateral flange or flaplike extension. Another specimen (UPNG

Vol. 6, p. 146

Asiatic Herpetological Research

TABLE 8. Sympatric female A. kunuaensis and A. solomonis scale counts
Species

kunuaensis

solomonis
kunuaensis

solomonis

locality

n

TSR

DC

June 1995

'

AMPSR

June 1995

Asiatic Herpetological ResearchVol. 6, p. 147

higher in A.solomonis than in A
kunuaensis at all sympatric localities. All

In Acutotyphlops infralabialis, MCZ
72129 has the prefrontal, preocular andthree Bougainville species of Acutotyphlops

anterior ocular partially fused into a single

are sympatric at Buin; unfortunately, the A.

10155 has the third andshield;fourth supralabials fused on the right side.

composed entirely of

infralabialis sample ismales.

Nevertheless, the three species (A.

infralabialis, A. solomonis, and A.

kunuaensis, respectively) are easily
distinguishable on head shape, headscutellation, coloration, and longitudinal=scale rows (MSR = 28, 32, 30;

AMPSR

90, 92-94, 95), while A. solomonis (x =exhibits significantly fewer
middorsals than either A. kunuaensis (525)or A. infralabialis (x = 508). Because the364.8)

three species retain their integrity throughout

and in areas of* sympatry,without any evidence of hybridization, they
their ranges

are justifiably recognized as valid species.

Anomalies of scutellation.

— In

Acutotyphlops subocularis, Hediger (1934)
and McDowell (1974) reported a specimen
lacking supranasals (NMBA 11704).

NMBA

and the supraocular fused to the prefrontal
on the left; and AMS 71360 exhibits one
supranasal on the right side.
In

Acutotyphlops kunuaensis,

fivesupralabials are present in two individuals(USNM 120936 and both sides of MCZ
65990), a T-V SIP in five specimens (bothsides of
72067, 77295; right side only

MCZ
MCZ 76957, 77003, 77298), and a T-VI
SIP on both sides of MCZ 77036; two
prefrontals are present in MCZ 72133,
76719, two frontals are present in MCZ
in76994, a suprarostral in three specimens(MCZ 76694, 76714, 76994), and paired
supraoculars in three specimens (both sidesof MCZ 76994, 77006; right side only in
MCZ 76697). Fusion of the preocular andthe other characters of this female are withinanterior ocular occurs in six individuals
76693, 76708, 76710,(both sides of77038; left side only of
76969; rightthe range of variation of A. subocularis. In

side only innumber of

preocular and anterior subocular occurs on
the left side of
76683, and fusion of
the preocular and postnasal occurs on the

Although they are absent bilaterally, all ofanterior

scale

rows

(40),

subcaudals (14), and suboculars (2 + 2),
plus the presence of a single shield between
nasal and ocular, it differs from A
solomonis so there can be no doubt about its
1 1709 has theidentity.
supraocular
fused to the ocular on the left side and11707 has the postocular fused tothe superior posterior subocular on the right
.

NMBA

NMBA
side.

AMS 41254 has five supralabials on

the right side (resulting

SL3);

PNGM

24601

from division of
exhibits

five

supralabials on both sides of the head (fromdivision of SL4); and24603 has the

PNGM

frontal divided into

two shields plus fivesupralabials on each side of the head (fromdivision of SL3).

NMV

In Acutotyphlops solomonis,
10108 has five supralabials on both sides,with the second to fourth occupying
positions of typical second and third; MCZ
65992, 65998, and 72138-39 have fivesupralabials on both sides; and MCZ 72138
has both the nasal and prefrontal
semidivided on the left side.

MCZ

MCZ
MCZ 76683), fusion of theMCZ

MCZ 76686. The prefrontal issemidivided in three specimens (left sideright side in

MCZ 72133, 76697; right side only
MCZ 76719) and the preocular andonly ininanterior ocular are each semidivided on bothsides in

MCZ 64236.

The preocular is

MCZ

divided on the right side only in
76961
while the preocular and anterior ocular are

semifused in two specimens (left side only
MCZ 77267 and 121909). Due to an

in

MCZ

175085 exhibits aapparent injury,
concavity and lacks a nostril on the rightside in addition to possessing extra scales inthat region. Based upon a sample of 510(counting condition on each side of the headseparately), the above cephalic scutellation

anomalies occur in frequencies of 0.2-1.6%
and may therefore be considered of rareoccurrence.
However, the increased
aberrations may be related to the conditions
that led to the original fragmentation of headshields within the group.

Laszlo Meszoly illustrated the holotypes ofSummary

A newgenus, Acutotyphlops
proposed to contain four species of highly
derived blind snakes from McDowell's(1974) Ramphotyphlops subocularis species
group (with the exclusion of/?, willeyi anditstransfer

June 1995

Asiatic Herpetological ResearchVol. 6, p. 148

,

to

the

is

Ramphotyphlops

flaviventer species group).

In addition to

Acutotyphlops subocularis (with its
synonym T. keasti), two species are revived
from synonymy {Acutotyphlops solomonisand Acutotyphlops infralabialis, the latterwith its synonyms T. bergi and T. adamsi)
and a new endemic species is describedTyphlops subocularis, T. infralabialis, T.
(Fig. 2), andkunuaensis
Acutotyphlops(Fig. 4). Thanks
are due to Roxane Coombs for executing theillustrations in Fig. 1 and proofreading themanuscript and to Kevin Wong fortransverse scale counts of A. kunuaensis
and statistical analyses of some of the data.
Charles Meszoely and the Biology
Department of Northeastern University

solomonis, T. adamsi

provided funding for the illustrations of thetype specimens.

Material ExaminedfromBougainville {Acutotyphlops
kunuaensis). Unique characters for thegroup include a parietal spike between thefrontal bones, multiple ocular, preocular
and/or subocular shields, a V-shaped lower
jaw, five or more infralabials, and afrontorostral shield with paired prefrontals.

Acutotyphlops is also known for its wedgeshaped or pointed head, a high number of
longitudinal scale rows, a multicameral right
lung, a straight unsegmented liver, lack of arectalceacum, Ramphotyphlops-\ike

hemipenis with 3-9 coils in retracted organ,and prominent sexual dimorphism in tailproportions and caudal counts. Five speciesof typhlopids are now known from
Bougainville Island: Acutotyphlops
infralabialis, A. kunuaensis, A. solomonis,

Ramphotyphlops braminus, and R.

flaviventer {= R. depressus).

Acknowledgements

Acutotyphlops kunuaensis (excluding
listedmaterial
above).
type—

BOUGAINVILLE IS.: No specific locality:

USNM 120211; Buin (6°50'S, 155°44'E,
ca. 60 m), MCZ 65990; Cape Torokina(6°15'S, 155°02'E, NSL), USNM 120949;
Empress Augusta Bay (6°25'S, 155°05'E,
NSL), FMNH 44800-01; Kieta (6°13'S,155°38'E, NSL), AMNH 87360-62; MCZ

NMV10109; Melelup
64226-36, 72104-05;(5°37'S, 154°55'E, ca. 915 m),
175089; Mutuhai (5°38'S, 154°57'E, ca.
820 m), MCZ 87605, 174754-55, 174759;
North Nasioi (ca. 6°10'S, 155°30'E),
6601014; Topanos (5°38'S, 155°00'E, 150
87606-07, 88049, 175082-88;m),
Torokina Bay (6°14'S, 155°03'E, NSL),
120931, 120933-34, 120935-48;
Torokina: Piva (6°14'S, 155°03'E, NSL),

MCZ
MCZ

MCZ

USNM

AMS 121582-84, 121698-700, 121769,
121909, 121956-57, 123393-99, 123402MCZ 76699.

03. Skull:

Appreciation is due to the curators andfollowing institutions wholoaned material for this study: Charles W.
staff of theMyers (AMNH), Alan E. Greer (AMS),
Carla H. Kishinami (BPBM), Jack W.
Sites, Jr. (BYU), Harold K. Voris, Robert
F. Inger, and Alan Resetar (FMNH), J.
Govaere and Mathias Lang (IRSNB), Jose
P. Rosado and Ernest E. Williams (MCZ),
Harry W. Greene (MVZ), Eugen Kramer(NMBA), A. J. Coventry (NMV), Ilaiah
Bigilale (PNGM), Arnold G. Kluge andGreg Schneider (UMMZ), James I. Menzies(UPNG), W. Ronald Heyer and Addison
H. Wynn (USNM), Rainer Gunther (ZMB),
and Hans-Wilhelm Koepcke (ZMH).

Acutotyphlops infralabialis.

—

BOUGAINVILLE:

Malabita (6°46'S,
65991, Buin,
155°43'E, ca. 150 m),
72129, Turiboiru (6°44'S, 155°41'E,
ca. 50 m),
92504; GUADALCANAL:
7040; Visale (9°15'S, 159°42'E),
71360, Mt. Austen (9°29'S,
77116; Makaruka
159°59'E, 245 m),
110249;(9°30'S, 160°04'E, 60 m),
Nalimbu River, 1 mi. inland (ca. 9°24'S,
40753 (holotype of
160°09'E, 15 m),

MCZ

MCZ

MCZ

BYU

AMS

AMS

MCZ

MVZ

T. adamsi);

MALAITA: AMS 4609

(holotype of T. infralabialis), vie. of(8°24'S, 160°36'E),
87396; Buma (8°56'S, 160°47'E),

Mbita'ama

AMS
NMBA

NEW

GEORGIA: Segi Point,
10155;
Horseshoe Reservation (8°34'S, 157°55'E),
95445 (holotype of T. bergi).
Skull: MCZ 64226.

UMMZ

Acutotyphlops solomonis.

—

BOUGAIN-VILLE: IRSNB 2029 (holotypeof T. solomonis); Buin, AMS 11451-52,
65999, 72084; Empress Augusta Bay,64225, 6599244802; Kieta,

MCZ
FMNH
98,

MCZ

New Britain, New Guinea, Loyalty Islands andelsewhere collected during the years 1895, 1896 and1897. Vol.5. Cambridge University, Cambridge.

BRONGERSMA, L. D.

1934.

Contributions toherpetology.
Mededeelingen 17(34):161-251.

Zoologische

Indo-Australian

1979. Type specimens of reptiles
and amphibians in the Australian Museum. Records

COGGER, H. G.

of the Australian Museum 32(4): 163-210.

NMV 10108; Kunua, MCZ 72083,

72085-86,72938,

72138-39,

73766,

76688, 175099; Melelup, MCZ 175090;
Mutuhai, MCZ 174756-58, 174760;
120932, 120934.
Torokina,

USNM

1944.

The skeletal

anatomy of Liotyphlops albirostris.
Morphology 74(2):287-295.

Journal of

DUNN, E. R., and J. A. TIHEN.

1976.
Aspects of the biology of
In A. d'A.
snakes.
Pp. 191-204.
BELLAIRS and C. B. COX (eds.), Morphology andbiology of reptiles. Linnean Society Symposium

PAPUA NEW GUINEA: Alotau (10°18'S,

GANS, C.

145955.
150°25'E, NSL),
72084.
65597, 65993,

uropeltid

MCZ

Skulls:

MCZ

—

PAPUA
Acutotyphlops subocularis.
Bismarck
GUINEA:
Archipelago:
IRELAND:ZMB 38612, 50458;

NEW

NEW

NMBA

,

Fissoa (2°55 S, 151°27'E, NSL),
11709-10; Lemkamin (3°20'S, 151°55'E),
5269; Medina (2° 54'S, 151° 22'E,

ZMUC

UPNG 5652; Radina (? =AMS41253-54; Yalom (4°25'S,
Medina),<100 m),ZMUC 5265-68;

1000 m),

151°45'E,

DUKE OF YORK IS. (4°10'S, 152°28'E,

AMS 2202 (holotype of T.
subocularis); NEW BRITAIN: Iambon, S<50 m),Whiteman Range

slope

(ca.

UPNG

1065

m),

1101;

152°16'E, NSL),

Kokopo(4°21'S,ZMH 3968; Mosa, West

Nakanai (5° 38'S, 150° 17'E, ca. 50 m),
PNGM 24600-03; Talasea (5°17'S,
150°02'E, NSL), MCZ 175091; Wunung,Jacquinot Bay (5°37'S, 151°27'E, NSL),NMBA 11705-08; UMBOI IS.: Awelkon(5°38'S, 147°50'E, 600 m), BPBM 5457;"New Guinea," ZMB 24341. Skull: NMBA
11704.

Unexamined

literature

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snakes: the freight-train approach to burrowing.
Science 199:189-192.
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und Reptilien: Anomalepididae, Leptotyphlopidae,
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HAHN, D. E.

KINGHORN, J. R.

1948.

Typhlops from New

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Records of the

LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, andC. E. DAWSON.

Standards in herpetology

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MCCOY, M.

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Reptiles of theSolomonIslands. Wau Ecology Institute Handbook (7): 1-80.

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