Breviora

B RE V I O R A

us ISSN 0006-9698

Cambridge, Mass. 7 January 2004 Number 512

TYPHLOPS LAZELLI, A NEW SPECIES OF CHINESE

BLINDSNAKE FROM HONG KONG

(SERPENTES: TYPHLOPIDAE)

V. Wallach^ and Olivier S. G. Pauwels^

Abstract. A new species of blindsnake is described from Hong Kong, China,
bringing the total number of scolecophidian species there to three and the number
of endemic snakes to three. This species is characterized by having 18 scale rows,
a T-V supralabial imbrication pattern, and a unicameral tracheal lung, and it ap-
pears to be a member of the Typhlops porrectus species group of South and
Southeast Asia.

INTRODUCTION

The scolecophidian fauna of China, including Hong Kong and
Taiwan, presently consists of four species placed in the genera
Typhlops and Ramphotyphlops. Although Typhlops diardii Schle-
gel (1839) and T. koshunensis Oshima (1916) are restricted to
southern China and Taiwan, respectively, Ramphotyphlops albi-
ceps (Boulenger, 1898) and R. braminus (Daudin, 1803) both are
known from Hong Kong (Karsen et al, 1986; Zhao and Adler,
1993; Zhao et al, 1998).

Typhlops (= Ramphotyphlops) braminus was first reported "on
the Peak in Hongkong Island" by Wall (1903) and for many years
was the only typhlopid known from there (Herklots, 1951). The

' Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cam-
bridge, Massachusetts 02138, U.S.A.; e-mail: vwallach@oeb.harvard.edu.
- Institut Royal des Sciences Naturelles de Belgique, Rue Vautier 29, 1000 Brus-
sels, Belgium; e-mail: osgpauwels@hotmail.com.

MCZ
LIBRARY

MAR 0 4 2004

HARVARD
I INIVERSITY

B RE V I O R A

us ISSN 0006-9698

Cambridge, Mass. 7 January 2004 Number 512

TYPHLOPS LAZELLI, A NEW SPECIES OF CHINESE

BLINDSNAKE FROM HONG KONG

(SERPENTES: TYPHLOPIDAE)

V. Wallach' and Olivier S. G. Pauwels-

Abstract. a new species of blindsnake is described from Hong Kong, China,
bringing the total number of scolecophidian species there to three and the number
of endemic snakes to three. This species is characterized by having 18 scale rows,
a T-V supralabial imbrication pattern, and a unicameral tracheal lung, and it ap-
pears to be a member of the Typhlops porrectus species group of South and
Southeast Asia.

INTRODUCTION

The scolecophidian fauna of China, including Hong Kong and
Taiwan, presently consists of four species placed in the genera
Typhlops and Ramphotyphlops. Although Typhlops diardii Schle-
gel (1839) and T. koshunensis Oshima (1916) are restricted to
southern China and Taiwan, respectively, Ramphotyphlops albi-
ceps (Boulenger, 1898) and R. braminus (Daudin, 1803) both are
known from Hong Kong (Karsen et al, 1986; Zhao and Adler,
1993; Zhao et al, 1998).

Typhlops {— Ramphotyphlops) braminus was first reported "on
the Peak in Hongkong Island" by Wall (1903) and for many years
was the only typhlopid known from there (Herklots, 1951). The

' Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cam-
bridge, Massachusetts 02138, U.S.A.; e-mail: vwalIach@oeb.harvard.edu.
- Institut Royal des Sciences Naturelles de Belgique. Rue Vautier 29. 1000 Brus-
sels, Belgium; e-mail: osgpauwels@hotmail.com.

MCZ
LIBRARY

MAR 0 4 2004

2 BREVIORA No. 512

existence of/?, albiceps in Hong Kong was verified in 1952 based
upon a specimen (BMNH 1954.1.13.4) originating from Caine
Road, Hong Kong Island, as reported by Romer (1970), who also
noted a second specimen (BMNH 1983.946) obtained from Car-
oline Hill Road, Hong Kong Island, in 1966. Taylor (1965) dis-
cussed a British Museum specimen (BMNH 1954.1.13.4) from
Hong Kong that was referred to R. albiceps but suggested that
its identity might be in error because its middorsal scales were
411 as compared with the then-known range of 301-327 in R.
albiceps. However, a range of 307-424 middorsals has been con-
firmed for the species (Wallach, 1998b). Hahn (1980) later ques-
tioned the species' presence in Hong Kong, presumably based on
Taylor's report. Karsen et al (1986) listed R. albiceps for the
fauna of Hong Kong based upon the two above-mentioned spec-
imens. Lazell and Lu (1990) referred two additional Hong Kong
specimens to Typhlops (= Ramphotyphlops) albiceps. These
specimens were collected by Sandra Brown (MackUn, 1988), with
one deposited at the St. Louis School (SLS 196), West Point,
Hong Kong, and the other at the Museum of Comparative Zo-
ology, Harvard University (MCZ 173290). Another specimen was
collected on the Hong Kong University campus by Michael Lau
in 1992; it also was deposited at the MCZ (MCZ 183578). In the
most recent herpetofaunal synopsis of Hong Kong, Karsen et al.
(1998) listed Hong Kong records of R. albiceps based upon five
specimens, including the British Museum pair discussed above
plus three that were collected in 1988 "in a patch of woodland
on the slope of Mt. High West." Two of the latter represent the
new species described below.

The known range of R. albiceps is southern Myanmar, southern
Thailand, peninsular Malaysia, the Kedah and Jarak Islands in the
Strait of Malacca, the Similan Islands off peninsular Thailand,
and the disjunct population from Hong Kong, China (Hahn, 1980;
McDiarmid et al, 1999). Zhao and Adler (1993) erroneously re-
ported R. albiceps from Singapore, presumably based upon Gran-
dison (1978), but this has not been verified by Lim and Chou
(1990) or Lim and Lim (1992).

All but two of the above specimens have been confirmed to
represent R. albiceps. However, the two MCZ specimens are not

2004 NEW HONG KONG TYPHLOPS 3

referrable to R. albiceps but instead represent an undescribed spe-
cies.

MATERIALS AND METHODS

All specimens were examined under a binocular microscope;
internal and external measurements were made to the nearest 0.5
mm with either vernier calipers or a metric ruler. Middorsal scales
were counted between the rostral scale and the terminal cone.
Dorsocaudals are defined as the number of vertebral scales along
the tail, counted between an imaginary line perpendicular to the
vent and the apical spine. The dorsocaudal count in samples usu-
ally is less variable than the count of midventral subcaudals,
which often are irregularly arranged.

Visceral characters have been defined and discussed in Wallach
(1985, 1993a, 1994, 1996, 1998a,b), Broadley and Wallach
(1996), and Wallach and Ineich (1996). All values of organ
lengths, gaps, intervals, and segments are given as percent snout-
vent length (% SVL), followed only by the % sign. Organ
lengths, measured as the maximum anterior-posterior distance,
are followed parenthetically by the organ midpoint (MP) value,
also as % SVL. Organ gaps are defined as the distance between
two organs (posterior tip of cranial organ to anterior tip of caudal
organ); organ intervals are defined as the distance between two
organs including the length of both organs (anterior tip of cranial
organ to posterior tip of caudal organ). Organ midpoint segments
are defined as the distance from the midpoint of one organ to the
midpoint of another organ.

The supralabial imbrication patterns (SIPs) of the Typhlopidae
consist of five states, each of which is denoted by the supralabial
numbers that overlap the shields dorsal to them: T-I with first
supralabial overlapping preocular, T-Il with second supralabial
overlapping preocular or presubocular, T-III with third supralabial
overlapping ocular or subocular, T-V with both second and third
supralabials overlapping shields above them, and T-0 with no
overlapping supralabials (Wallach, 1993b).

Museum acronyms follow Leviton et al. (1985), except for the
following: SLS = St. Louis School, Hong Kong; TNRC = Thai-

BREVIORA

No. 512

Figure 1. Head of the holotype of Typhlops lazelli (MCZ 173290). A. dorsal
view; B, lateral view.

land National Reference Collection. Bangkok: ZRC = Zoological
Reference Collection, Singapore National University, Singapore.

TAXONOMY

Typhlops lazelli, new species
Figure 1

Holotype. MCZ 173290, adult female collected by Sandra
Brown (field no. Z-36266) on 27 May 1988.

Type Locality. High West, Pokfulam, Hong Kong Island, Hong

2004 NEW HONG KONG TYPHLOPS 5

Kong Territory, China, ca. 22°15'30"N, 114°08'30"E (mapped by
Lazell and Lu, 1990: fig. 1, locality 3).

Paratype. MCZ 183578 (ex-SLS), a juvenile male from Hong
Kong University Campus, Hong Kong, Hong Kong Territory,
China, collected by Michael Wai-Neng Lau on 10 December
1992.

Diagnosis. Typhlops lazelli can be distinguished from all other
typhlopids of Asia except T. porrectus by the combination of 18
midbody scale rows and a T-V SIP. From T. porrectus, it is dif-
ferentiated by a unicameral tracheal lung, absence of enlarged
occipitals, and a projecting mental shield.

Etymology. This species is dedicated to James D. "Skip" La-
zell in recognition of his exemplary studies of the herpetofauna
of the Hong Kong environs. As one of the vanishing breed of
19th century naturalists, Skip is an arduous field worker, an ex-
emplary systematist, and an expert on island biogeography. His
many contributions to Caribbean and Chinese insular herpetology
attest to his authority in the field. Although his writings may at
times be acerbic and contentious, they are always honest, insight-
ful, and entertaining.

Description of Holotype (Variation in Paratype Given Paren-
thetically). Adult female (juvenile male) with snout-vent length
of 155.2 (91.1) mm, tail length of 2.8 (1.4) mm, total length 158.0
(92.5) mm, midbody diameter 1.9 (1.2) mm, midtail diameter 1.7
(0.8) mm, total length/midbody diameter ratio 83.2 (77.1), tail
1.8% (1.5%) of total length, tail length/tail width ratio 1.65 (1.75).
Longitudinal scale rows 17-18-18 (17-18-18), total middorsals
427 (409), subcaudals 10 (9), dorsocaudals 9 (8). Three anal
scales. Apical spine short and straight with stout base.

Head rounded in dorsal view, not distinct from the neck, with
a truncated snout. Rostral oval, 0.32 (0.33) head diameter and
slightly broader than supranasals, not reaching the level of the
eyes, in contact with frontal that separates the supranasals mid-
dorsally. Frontal 2.0 (2.0) times as broad as long with rounded
posterior border, smaller than supraoculars. Supraoculars trans-
versely oriented, 1.5 times the size of costal or body scales. Pa-
rietals transverse, twice the size of the costals. Discretely enlarged
occipitals lacking, subequal to costals in size. Postfrontal, inter-

6 BREVIORA No. 512

parietal, and interoccipital larger and broader than frontal. Snout
rounded in lateral view, supranasal broader than preocular with
weak postnasal concavity. Nasal incompletely divided with in-
ferior nasal suture contacting second supralabial. Superior nasal
suture curving dorsomedially in an arc, extending 0.90 (0.67) of
the nostril-rostral gap and just visible in dorsal view. Nostril clos-
er to rostral than preocular, directed laterally, its axis oriented at
45°. Ocular subequal in size but slightly narrower than preocular.
Eye barely visible as a vague pigmented spot (discrete eyespot)
beneath the ocular-supraocular suture near the supraocular-pre-
cular junction. One postocular. Four supralabials, T-V SIP with
both the second and third supralabials overlapping the shields
above them. First supralabial half the size of second, second su-
pralabial two-thirds the size of third, and third supralabial one-
half the size of fourth. Mental not projecting beyond curvature
of lower jaw. Three scales border cloacal opening. Tail with
abrupt taper near tip. Apical spine lacking, tail terminus covered
by an obtusely pointed cone. An anomalous condition in the para-
type is the partial fusion, on both sides of the head, of the dor-
solateral portion of the postnasal with the preocular and the preo-
cular with the ocular.

Cephalic glands confined to sutures between scales. One pair
of lateral tongue papillae present just proximal to level of bifur-
cation of lingual tips.

Middorsal nine scale rows pigmented lightly brown with a
darker brown spot covering anterior V^-V^ of scale; midventral
nine rows lightly stippled in brown with white background and
outer margins. Anterior snout (most of rostral, nasals, peroculars,
oculars, and labials), chin, and throat white; a median longitudinal
white bar occurs on throat of holotype. Rostral of holotype white
with central brown bar; in paratype the entire rostral is brown.
Ventrally, the cloacal region to tail tip white. Paratype with a few
scattered midventral white scales.

Internal anatomy (female holotype data first and, if different,
male paratype data second). Sternohyoideus posterior tip 7.4,
9.3%, sternohyoideus-heart gap 0.74, 0.71. Heart 3.2, 3.6% (MP
30.0, 33.5%), systemic arch gap/heart length ratio 0.20, 0.15.
Snout-heart interval 31.6, 35.2%, heart-liver gap absent (0,

2004 NEW HONG KONG TYPHLOPS 7

-0.3%) with anterior tip of left liver lobe just touching (overlap-
ping) the posterior tip of ventricle. Liver tightly coiled, 19.4,
25.7% (MP 45.2, 50.5%), left anterior lobe extends beyond right
by 0.10, 0.17 liver length, right posterior tail extends beyond left
by 0.44, 0.47 liver length, left liver/right liver ratio 0.63, 0.61.
Right liver segments 17, 15, left liver segments 12, 10, total liver
segments 29, 25. Liver-gall bladder gap 7.7, 9.3%, liver-gall
bladder interval 35.8, 28.4%. Gall bladder 1.3, 0.8% (midpoint
63.2, 73.1%) located between spleen and pancreas, spleen (1.0%)
craniad of and separated from the pancreas (1.0%). Right ovary
2.3% (MP 70.8%) with 6 follicles, left ovary 2.3% (MP 75.6%)
with 4 follicles. Testes unipartite, right testis 2.2% (MP 76.5%),
left testis 2.2% (MP 79.8%). Right adrenal MP 85.1, 85.9%, left
adrenal MP 86.7, 87.8%. Liver-kidney gap, 28.4, 21.9%, liver-
kidney interval 58.7, 57.4%. Kidneys not segmented but with
dark striations, each with a single renal artery, right kidney 3.9,
3.6% (MP 85.2, 87.0%), left kidney 3.9, 3.3% (MP 88.4, 90.7%),
kidney-vent interval 16.8, 14.8%, kidney-vent gap 9.7, 7.7%.
Elongate rectal caecum present (1.6, 0.8%), twice the diameter
of adjacent intestine, caecum-vent interval 10.0, 6.3%. Caecum/
left kidney ratio 0.41, 0.24.

The tracheal lung, cardiac lung, and right lung are unicameral;
tracheal organ with 34 transverse septa forming incipient pauci-
cameral cells, each with septa twice the height of the faveoli. Left
lung complex absent. Trachea 3L0, 34.2% (MP 16.1, 18.2%) pos-
sessing short tips on cartilaginous rings, numbering an estimated
240, 279 rings (rings/10% SVL = 77.5, 82.4). Tracheal lung 19.3,
25.1% (MP 18.7, 19.1%), saccular with 34 transverse blood ves-
sels serving it. Anterior tip of parenchyma 9.0, 6.6%; posterior
tip of parenchyma 40.6, 41.5%. Tracheal membrane/trachea ratio
large, 3.0 posteriorly, 4.0 at midneck, and 1.0 anteriorly. Right
lung 18.7, 161.% (MP 41.0, 43.3%), poorly vascularized with
very large ediculae in cranial portion, caudal portion (9.7, 9.8%)
with large trabeculae, posterior lung tip at 50.3, 51.4%. Right
bronchus 9.0, 10.1%, bronchus/right lung ratio 0.48, 0.63, right
lung/tracheal lung ratio 0.96, 0.64, total lung 41.3, 44.8% (MP
29.0, 29.7%).

Organ midpoint segments include heart-right lung segment

8 BREVIORA No. 512

(11.0, 9.8%), heart-liver segment (15.2, 17.0%), trachea-liver
segment (29.1, 32.3%), heart-right lung segment (40.8, 43.0%),
liver-kidney segment (41.6, 38.4%), trachea-bronchus — gall
bladder segment (42.6, 49.9%), right lung-adrenal segment (44.9,
43.6%), heart-kidney segment (56.8, 55.4%), trachea-bronchus —
kidney segment (66.2, 65.7%), and trachea-adrenal segment
(69.8, 68.7%).

Ecology. The only specimens found thus far were removed
from leaf litter in a concrete drainage ditch into which they were
presumably washed by rain from the forested slopes above the
drain. Most upland areas of Hong Kong Island are covered in
richly wooded forest today, often with large trees (Herklots,
1951). These areas have been protected as a country park to in-
sure sufficient watershed for the island's reservoirs (J. D. Lazell,
personal communication).

It is perhaps surprising that T. lazelli has not been found on
any other of the more than 100 islands of Hong Kong, or on the
New Territories mainland. However, few areas exist anywhere in
tropical China today as well forested as the uplands of Hong
Kong Island (J. D. Lazell, personal communication). Indeed, py-
thons, cobras, and ratsnakes, in addition to many smaller species
of snakes, are common on the island (Karsen et aL, 1986). If the
forest is the natural home of T. lazelli, then its future would
appear to be secure. Our ability to obtain further specimens will
depend upon luck and the cooperation of local residents in search-
ing drain gutters.

DISCUSSION

All seven cuiTently recognized typhlopid genera are found in
the Old World (Acittonphlops, Cyclotyphlops, Ramphotyphlops,
Rhinotyphlops, Typhlops, Xenotyphlops, and Giypotyphlops Pe-
ters, 1881 [resurrected for "'Rhinotyphlops'' acutus by Wallach,
2003]), with six occupying or extending into Asia. Among the
150 species of Typhlopidae examined thus far, no species of Acu-
totyphlops, Cyclotyphlops, Grypotyphlops, Ramphotyphlops,
Rhinotyphlops, or Xenotyphlops has a unicameral tracheal lung
but nine species of Typhlops possess it: two African (T. caecatiis
and T. zenkeri), three Asian (T. depressiceps, T. hedraeus, and T.

2004 NEW HONG KONG TYPHLOPS 9

minis), and four American (T. monensis, T. pusiUus, T. rostella-
tus, and T. scJnvartzi) (Wallach, 1998b).

The T-V SIP of T. lazelli is consistent only with the genus
Typhlops. The other six genera are separable on the following
characters: Aciitotyphlops has a T-0 or T-III SIP, 26-36 midbody
scale rows, fragmentation of head shields, and a pointed snout;
Cycloty'phlops has a T-III SIR 22 scale rows, and a floral pattern
of head scales; Grypotyphlops has a T-0 SIP and 24-34 scale
rows; Ramphotyphlops has a T-III SIP; Rhinotyphlops has a T-0
or T-II SIP; and Xenotyphlops has a T-0 SIP Typhlops may have
any of the possible imbrication patterns (T-0, T-II, T-III, or T-V).

It was previously believed that presence of lateral tongue pa-
pillae was a diagnostic feature of certain Ramphotyphlops. Mc-
Dowell (1974) listed 10 species oi Ramphotyphlops and only one
of Typhlops {Typhlops {— Giyponphlops] aciitiis) that possess
lingual papillae (plus a few Ramphotyphlops species that either
lacked papillae or were variable). Wallach (1998b) reported pa-
pillae in three additional Ramphotyphlops, two Rhinotyphlops,
and 12 additional Typhlops. Thus, the presence of papillae is not
confined to a particular genus and is uninformative.

The T-V SIP and unicameral tracheal lung are compelling ev-
idence for placement in Typhlops versus RamphotypJilops. Ty-
phlops lazelli appears to be allied to the Typhlops porrectus spe-
cies group of southern and eastern Asia. However, a comparison
with all Asian species groups in these two genera follows.

1. The Ramphotyphlops braminus species group contains only
R. bramimts (Daudin, 1803), the parthenogenetic "flowerpot"
species that is distributed worldwide. It differs from T. lazelli in
having 20 scale rows, superior nasal suture extending onto dor-
sum of snout, inferior nasal suture contacting preocular, and a
T-III SIR

2. The Ramphotyphlops lineatiis species group is also mono-
typic, with R. lineatus (Schlegel, 1839) as its sole member. It
differs from T lazelli in having 22-24 scale rows, a T-III SIP,
inferior nasal suture contacting first supralabial, striped color pat-
tern, and in lacking a visible eye.

3. Typhlops marxi Wallach (1994), cuiTently a monotypic
member of the T marxi species group, may actually belong to

10 BREVIORA No. 512

the long-tailed Rciniphotyphlops iniiltilineatiis species group: R.
cumingii (Gray, 1845), R. multilineatus (Schlegel, 1839), and R.
olivaceiis (Gray, 1845). Two of the species are Philippine, and
other than the T-0 SIP, T. marxi agrees most closely with R.
cumingii in having a high number of midbody scale rows (30),
total middorsals (525), and subcaudals (36), in addition to a
keeled rostral and relatively long tail (6% total length) (Wallach,
1994). If so included, T. marxi would be the only Ramphotyph-
lops with a T-0 SIP; however, an identical situation is seen in
Acutotyphlops, where three of the four species — A. infralabialis
(Waite, 1918), A. kunuaensis Wallach, 1995, and A. solomonis
(Parker, 1939) — have a T-III pattern, with A. subocularis (Waite,
1897) having a T-0 SIP Therefore, we suggest the transfer of T.
marxi to the R. multilineatus species group as Ramphotyphlops
marxi (Wallach, 1993a) comb. nov. Members of this group can
be separated from Typhlops lazelli by their midbody scale rows
(20-30), long tails (3-9% total length), and pointed snouts with
angular rostral edges.

4. The Typhlops ater species group {sensu Wallach, 1996) is
characterized by cephalic glands distributed beneath the central
regions of head shields (in addition to peripherally, as in all other
typhlopids), a T-II or T-V SIP, broad rostral, nasal usually divid-
ed, and absence of a rectal caecum. This species group contains
the following 15 species: T. andamanensis Stoliczka, 1871; T.
ater Schlegel, 1839; T. beddomii Boulenger, 1890; T. bisubocu-
laris Boettger, 1893; T. ceylonicus Smith, 1943; T. depressiceps
Sternfeld, 1913; T. floweri Boulenger in Flower, 1899; T. fred-
parkeri Wallach, 1997; T. hedraeus Savage, 1950; T. inornatus
Boulenger, 1888; T. mcdowelli WaWsich, 1997; T. mirus Jan. 1863;
T. oligolepis Wall, 1909a; T. thurstoni Boettger, 1890; and T.
tindalli Smith, 1943. Most mainland Asian species have 18 scale
rows, whereas species from the East Indies have from 16 to 24
midbody rows.

In addition to the above characters of the T. ater group that
distinguish all species from T. lazelli, the following species with
a T-V SIP can be further differentiated as follows: T. depressiceps
has 20-24 midbody scale rows, more than 630 middorsals, more
than 20 subcaudals, and a hooked snout; T. floweri has more than

2004 NEW HONG KONG TYPHLOPS U

475 middorsals, a suboculai; 20 or more subcaudals, and a mul-
ticameral lung; T. inomatus has 20-22 scale rows and a subcular;
and T. mcdowelli has 22-24 scale rows, 17 or more subcaudals,
and a rostral beak.

5. The Typhlops diardii {sensu McDowell, 1974; Wallach,
2001 ) species group, characterized by 20-30 midbody scale rows,
a T-V SIP, broad rostral, nasal incompletely divided, large pe-
dunculate rectal caecum, and a short tail that is broader than long,
contains 11 species: T. bothhorhynchiis Gunther, 1864; T. diardii
Schlegel, 1839; T. giodinhensis BouiTCt, 1937; T. hypsobothriiis
Werner, 1917; T. klemmeri Taylor, 1962; T. koshunensis Oshima,
1916; T. mueUeri Schlegel, 1839; T. oatesii Boulenger, 1890; T.
roxaneae Wallach, 2001; T. siamensis Gunther, 1864; and T. tran-
gensis Taylor, 1962. All of these species are clearly separable
from T. lazelli.

6. The primarily Philippine Typhlops ruficaiidus species group
{sensu McDowell, 1974), with a strongly contrasting bicolor pat-
tern (dark dorsum and light venter), 24-30 midbody scale rows,
a T-III SIP, and vestigial or absent rectal caecum, contains seven
species: T. castanotiis Wynn and Leviton, 1993; T. coUaris Wynn
and Leviton, 1993; T. fusciis Dumeril and Bibron, 1844; T. krciali
Doria, 1874; T. rwZ^^r Boettger, 1897; T. nificaiidiis (Gray, 1845);
and T. siduensis Taylor, 1918. They all are easily distinguishable
from T. lazelli by the characters above.

7. The Typhlops pammeces species group (previously referred
to as the '^ Ramphoty phi ops' ^ bramimis species group by Wallach,
1993a) with 20 scale rows (22 in T. leucomelas). T-III SIP, nar-
row rostral, and divided nasal with superior nasal suture visible
dorsally, has 10 members: T. conradi Peters, 1874; T. jerdoni
Boulenger, 1890; T. khoratensis Taylor, 1962; T. lankaensis Tay-
lor, 1947; T. leucomelas Boulenger, 1890; T. malcolmi Taylor,
1947; T. pammeces Gunther, 1864; T. tenebrarum Taylor, 1947;
T. veddae Taylor, 1947; and T. violaceiis Taylor, 1947. The va-
lidity of some of Taylor's species is uncertain and A. H. Wynn
currently is studying them. All of these species are distinguishable
from T. lazelli by the above suite of characters. Two Rampho-
typhlops species, R. albiceps Boulenger, 1898, and an undescribed
form from Thailand, also have been associated with this group.

12 BREVIORA No. 512

Although superficially similar, both have the characteristic hem-
ipenes of Ramphotyphlops in addition to 20 scale rows, a T-III
SIP, and a nasal suture extending onto dorsum of snout. The hem-
ipenis type of Sulawesi Typhlops conradi is unknown and it may
be a Ramphotyphlops, because it closely resembles R. albiceps.

8. The remaining Asian assemblage is the Typhlops porrectus
species group {sensu Wallach, 1999), characterized by 18 scale
rows, a narrow rostral, and a tail that is longer than broad. All
examined species except T. porrectus have a paucicameral tra-
cheal lung. It contains at least six species: T. exiguus Jan, 1864;
T. filiformis Dumeril and Bibron. 1844; T. loveridgei Constable,
1949; T. meszoelyi Wallach, 1999; T. porrectus StoHczka, 1871;
and T. schnmtzi Auffenberg. 1980. A number of uncertain names
{T. ahsanuli Khan. 1999b; T. ductuliformes Khan, 1999a; T.
mackinnoni Wall, 1909b; T. m. madgemintonae Khan, 1999b; T.
m. shermani Khan, 1999b; and T. venningi Wall, 1913) may or
may not be valid but appear to be related to T. porrectus (Wal-
lach, 1999, 2000). The group may be polyphyletic because it
includes species with T-II, T-III, and T-V SIPs, the SIP usually
diagnostic of species groups or even genera as previously men-
tioned. Work in progress by A. H. Wynn will hopefully establish
the content and relationships of the species in this group. Ty-
phlops lazelli is compared to other Asian Typhlops species having
18 scales and a T-V SIP in Table 1.

Typhlops lazelli can be separated from T. porrectus, which it
closely resembles in having 18 scale rows, a T-V SIR a narrow
rostral, an incomplete superior nasal suture, lateral tongue papil-
lae, and similar coloration, by its smoothly contoured mental
shield (vs. projecting) and lack of enlarged occipitals (vs. occip-
itals 1.5-2.0 times costal width). More significantly, T. lazelli also
can be separated from T porrectus by its unicameral tracheal
lung; T porrectus possesses a multicameral tracheal lung with
17-24 type C foramina. No evidence exists that the tracheal lung
structure changes ontogenetically such that a unicameral lung
transforms into a paucicameral and eventually into a multicameral
lung (Wallach, 1998b). Neonates and juveniles of large-sized spe-
cies that have multicameral lungs exhibit the same structure, in
miniature, as adults. Examples of such cases from three different

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genera include dissections of specimens with the following total
lengths: T. bibronii (117 mm vs. 356 mm), T. congestiis (204 mm
vs. 528 mm), T. jamaicensis (102 mm vs. 302 mm), T. lineolatiis
(120 mm vs. 302 mm), T. muelleri (116 mm vs. 369 mm), T.
punctatus (168 mm vs. 379 mm), T. nificaudus (127 mm vs. 367
mm), Ramphotyphlops albiceps (124 mm vs. 296 mm). Rain, ni-
grescens (140 mm vs. 281 mm). Ram. olivaceiis (163 mm vs.
358 mm), Rhinotyphlops mucruso (120 mm vs. 743 mm), and
Rhi. schlegelii (144 mm vs. 447 mm). All of these specimens
have multicameral tracheal lungs of the same form in both the
juveniles and adults. The difference between a unicameral and
multicameral lung is morphologically significant; no species has
yet been observed to exhibit more than a single lung type (uni-
cameral, paucicameral, or multicameral). There can thus be no
doubt about the validity of Typhlops lazelli.

The following keys will aid in the identification of Chinese and
Hong Kong Typhlopidae.

Key to Hong Kong Species of Typhlopidae

la. Inferior nasal suture contacting preocular. eye distinct with pupil, head brown

R. braminus

lb. Inferior nasal suture contacting second supralabial, eyespot indistinct, snout

white 2

2a. Supralabial imbrication pattern T-III, 20 scale rows, entire head and nape

white or yellow plus subcaudals and tail tip R. albiceps

2b. Supralabial imbrication pattern T-V, 18 scale rows, snout, chin, and throat

white plus subcaudals T. lazelli

Key to Chinese Species of Typhlopidae

la. Supralabial imbrication pattern T-III {Ramphonphlops) 2

lb. Supralabial imbrication pattern T-V (Typhlops) 3

2a. Scale rows 20, head brown, inferior nasal suture to preocular R. braminus
2b. Scale rows 18, head yellow or white, inferior nasal suture to second supraabial

R. albiceps

3a. Scale rows 18, middorsals greater than 400, length/width ratio greater than

60, one postocular T. lazelli

3b. Scale rows 22 or more, middorsals fewer than 350, length/width ratio less

than 50, two postoculars 4

4a. Scale rows 22 (23), middorsals fewer than 250 (Taiwan) . . . T. koshunensis
4b. Scale rows 24-28, middorsals greater than 260 (China) T. diardii

2004 NEW HONG KONG TYPHLOPS 15

Recent summaries of native Hong Kong reptiles and amphib-
ians reveal slightly different figures. Karsen et al. (1986, 1998)
listed 99 total species, whereas Lazell (1999) reported only 90
species, including five endemics. Of this number, Karsen et al.
(1986) listed 47 snakes, Karsen et al. (1998) listed 49 snakes,
and Lazell (1999) listed 38 snakes; all agreed that two species
are endemic. The presence of T. lazelli increases the number of
native Hong Kong snakes to either 39 or 50, with three endemic
species. Lazell (1999) hypothesized a Cathaysian origin for T.
koshimensis from Taiwan, a Sundaland origin for T. (= Rampho-
typhlops) albiceps of Southeast Asia, and unknown origins for T.
(= Ramphotyphlops) braminus and T. diardii. Ramphotyphlops
braminus, with many similar species in India and Southeast Asia,
would appear to have a western origin. Because T. diardii also
occurs throughout Southeast Asia, it could have a southern or
western origin. Typhlops lazelli, as a member of the T. porrectus
group, would have a western origin.

MATERIAL EXAMINED

Cyclotyphlops deharx^engi (MNHN 1990.4279 [holotype]),
Ramphotyphlops albiceps (BMNH 1946.1.10.50 [holotype],
1954.1.13.4, 1983.946; MCZ 181196, 177983; SLS 196; ZMUC
52203-04; ZRC 2.3043-45), R. ozakiae (FMNH 180003-06
[paratypes], 180007 [holotype]), Typhlops ate r {¥MNH 142108;
MCZ 33505; NMBA 979; ZMA 17737), T. beddomii (MCZ
3913, 3929, 22372, 175867; FMNH 217694), T. bisubocularis
(USNM 43455), T. bothriorhynchus (UF 48813), T. castanotus
(CAS 127973; CAS-SU 27942; MCZ 25594), T. ceyloniciis
(BMNH 1946.1.1 1.62 [holotype]), T. collaris (UF 54186, 68443),
T. conradi (ZMB 7934 [holotype]), T. depressiceps (MCZ
145954; USNM 195953; ZMB 23986 [holotype]), T. diardii
(CAS-SU 13982; FMNH 180008, 180023, 252064; MCZ
165004; ROM 25640), T. exigiius (ZMB 50030), T. filiformis
(MNHN 929 [holotype]), T. floweri (CAS 101599; MCZ 181 198;
NMBA 328), T. fredparkeri (MCZ 142651 [holotype]), T. fuscus
(MNHN 1062 [holotype]), T. hedraeus (MCZ 17578; USNM
229285, 498958), T. inoruatus (MCZ 140724, 140728, 175100;
UPNG 8572), T. jerdoni (ZMUC 52121), T. khoratensis (MCZ

16 BREVIORA No. 512

74097), T. klemmeri (FMNH 178238 [holotype]), T. kraalii
(ZMA 14225), T. lankaensis (FMNH 100134 [paratype]), T. leu-
comelas (BMNH 1946.1.10.46 [holotype]), T. lazelli (MCZ
173290 [holotype]; MCZ 183578 [paratype]), T. lovericlgei (MCZ
2283 [holotype]), T. malcolmi (FMNH 100132 [paratype]; ZMH
3967), T. marxi (FMNH 96520 [holotype]), T. mcdowelli (UPNG
7502 [holotype]; PNGM 24604 [paratype]), T. meszoelyi (FMNH
191888 [paratype], 191889 [holotype]), T. mirus (FMNH
123533-34; MCZ 18377-78; NHRM 3350), T. muelleri (BPBM
2156; CAS 222410; FMNH 161275, 252063; IRSNB 16535;
TNRC 3788, 7336-37; USNM 86885), T. pammeces (BMNH
1946.1.11.34 [holotype]; CM 90600; MCZ 5229; USNM
193298), T. porrectus (CAS 17169; FMNH 60645, 217449; MCZ
3702, 4082, 165023-24; NHRM 5529, UMMZ 123429 [paratype
of T. ductuliformes]), T. roxaneae (MCZ 177984 [holotype]), T.
ruber (CAS 182566; FMNH 53223; MCZ 79698), T. nificaudus
(CAS 135667; CAS-SU 19517, 21066, 26815; UF 54652), T.
schmutzi (UF 29452, 29528, 37018 [paratypes]), T. siamensis
(MCZ 16655; TCWC 29356), T. tenebrarum (FMNH 120237-38
[paratypes], 167012; NHRM 31445a), T. trangensis (FMNH
178236 [holotype]; NMV 1914), T. veddae (FMNH 100033 [ho-
lotype]), T. violaceiis (FMNH 100068 [holotype], 124231).

ACKNOWLEDGMENTS

The types of T. lazelli, collected by Sandra Brown and Michael
Lau Wai Neng, were donated to the MCZ by Anthony Bogadek.
Skip Lazell was responsible for making the specimens available
for study and arranging for their subsequent donation to the MCZ.
The illustrations of T. lazelli were drawn by the second author.
We wish to thank the following curators for permission to ex-
amine and dissect the comparative material that forms the basis
of this report; E. N. Arnold, B. Clarke, and C. J. McCarthy
(BMNH); C. Kishinami (BPBM); R. C. Drewes and J. V. Vindum
(CAS); C. J. McCoy, Jr., J. Wiens, and E. J. Censky (CM); H.
Marx, H. K. Voris, and R. F Inger (FMNH); E. E. Williams, J.
Hanken, and J. P Rosado (MCZ); I. Ineich (MNHN); S. O. Kul-
lander (NHRM); E. Kramer and B. Schatti (NMBA); A. J. Cov-
entry (NMV); I. Bigilale (PNGM); R. W. Murphy (ROM); A.

2004 NEW HONG KONG TYPHLOPS 17

Bogadek (SLS); J. R. Dixon (TCWC); W. Auffenberg and D. L.
Auth (UF); R. A. Nussbaum (UMMZ); J. Menzies (UPNG); G.
R. Zug and A. H. Wynn (USNM); L. van Tuijl (ZMA); R. Giin-
ther (ZMB); H.-W. Koepcke and J. Hallermann (ZMH); and J. B.
Rasmussen (ZMUC).

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