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us ISSN 0006-9698 

Cambridge, Mass. 7 January 2004 Number 512 




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. 


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: 
- Institut Royal des Sciences Naturelles de Belgique, Rue Vautier 29, 1000 Brus- 
sels, Belgium; e-mail: 


MAR 4 2004 



us ISSN 0006-9698 

Cambridge, Mass. 7 January 2004 Number 512 




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. 


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: 
- Institut Royal des Sciences Naturelles de Belgique. Rue Vautier 29. 1000 Brus- 
sels, Belgium; e-mail: 


MAR 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 


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


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- 


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. 


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 


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 

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, 


-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. 


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. 


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 

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 


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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14 BREVIORA No. 512 

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 


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. 


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). 


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. 


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). 


Auffenberg, W. 1980. The herpetofauna of Komodo, with notes on adjacent areas. 
Bulletin of the Florida State Museum (Biological Sciences), 25: 39-156. 

BoETTGER, O. 1890. Neue Schlange aus Ostindien. Berichte der Senckenbergisch- 
en Naturforschenden Gesellschaft in Frankfurt am Main, 1890: 297-298. 

. 1893. Neue Reptilien und Batrachier aus West-Java. Zoologischer An- 

zeiger, 16: 334-340. 

. 1897. Neue Reptilien und Batrachier von den Philippinen. Zoologischer 

Anzeiger, 20: 161-166. 

BouLENGER, G. A. 1888. Descriptions of new reptiles and batrachians obtained 
by Mr. H. O. Forbes in New Guinea. Annals and Magazine of Natural History, 
sen 6, 1: 343-346. 

. 1890. The Fauna of British India, Including Ceylon and Burma. Reptilia 

and Batrachia. London, Taylor and Francis. 541 pp. 

. 1898. Descriptions of two new blind snakes. Annals and Magazine of 

Natural History, sen 7, 1: 124. 

BouRRET, R. 1937. Notes herpetologiques sur ITndochine frangaise. XV. Lezards 
et serpents regus au Laboratoire des Sciences Naturelles de TUniversite au 
cours de I'annee 1937. Descriptions de deux especes et de deux varietes 
nouvelles. Bulletin General de ITnstruction Publique, Hanoi, 1937: 57-80. 

Broadley, D. G., and V. Wallach. 1996. A remarkable new worm snake (Ser- 
pentes: Leptotyphlopidae) from the East African coast. Copeia, 1996: 162- 

Constable, J. D. 1949. Reptiles from the Indian Peninsula in the Museum of 
Comparative Zoology. Bulletin of the Museum of Comparative Zoology. 103: 

Daudin, F M. 1803. Histoire Naturelle Generale et Particuliere des Reptiles. Tome 
Septieme. Paris, F Dufart. 436 pp. 

DoRiA, G. 1874. Enumerazione dei rettili raccolti dal Dott. O. Beccari in Amboina, 
alle Isole Aru ed alle Isole Kei durante gli anni 1872-73. Annali dell Museo 
Civico di Storia Naturale 'Giacomo Doria' di Genova, 6: 325-357. 

DuMERiL, A. M. C, AND G. BiBRON. 1844. Erpetologie Generale ou Histoire Na- 
turelle Complete des Reptiles. Tome Sixieme, Comprenant 1' Histoire Gener- 
ale des Ophidiens, la Description des Genres et des Especes de Serpent Non 
Venimeux, Savoir: La Totalite des Vermiformes ou des Scolecophides, et 
Partie des Circuriformes ou Azemiophides; en Tout Vingt-cinq Genres et 
Soixante-cinq Especes. Paris, Librairie Encyclopedique. 609 pp. 

Flower, S. S. 1899. Notes on a second collection of reptiles made in the Malay 

18 BREVIORA No. 512 

Peninsula and Siam, from November 1896 to September 1898, with a list of 
the species recorded from these countries. Proceedings of the Zoological So- 
ciety of London. 1899: 600-696. 

Grandison. a. G. C. 1978. Snakes of West Malaysia and Singapore. Annalen des 
Naturhistorischen Museums in Wien. 81: 283-303. 

Gray. J. E. 1845. Catalogue of the Specimens of Lizards in the Collection of the 
British Museum. London, British Museum (Natural History). 289 pp. 

GuNTHER, A. C. L. G. 1864. The Reptiles of British India. London. Robert Hard- 
wicke. 452 pp. 

Hahn. D. E. 1980. Liste der rezenten Amphibien und Reptilien: Anomalepididae, 
Leptotyphlopidae, Typhlopidae. Das Tierreich, 101: 1-93. 

Herklots, G. a. C. 1951. The Hong Kong Countryside. Hong Kong, South China 
Morning Post. 175 pp. 

Jan, G. 1863. Elenco Sistematico Degli Ofidi. Descritti e Disegnati per 
ITconografia Generale. Milan, Italy, A. Lombardi. 143 pp. 

. 1864 in G. Tan and F. Sordelli. 1860-1866. Iconographie Generale des 

Ophidiens. Tome Premiere (Livrais. 1 a 17) Contenant Cent Deux Planches. 
Milan, Italy, Georges Jan and Ferdinand Sordelli. 11 pp.. 102 pis. 

Karsen, S. J.. M. W.-N. Lau, and A. Bogadek. 1986. Hong Kong Amphibians 
and Reptiles. Hong Kong. Urban Council. 136 pp. 

. 1998. Hong Kong Amphibians and Reptiles. 2nd edition. Hong Kong, 

Provisional Urban Council. 186 pp. 

Khan, M. S. 1999a. Typhlops diictuliformes a new species of blind snakes from 
Pakistan and a note on T. porrectus Stoliczka, 1871 (Squamata: Serpentes: 
Scolecophidia). Pakistan Journal of Zoology, 31: 385-390. 

. 1999b. Two new species and a subspecies of blind snakes of genus 

Typhlops from Azad Kashmir and Punjab, Pakistan (Serpentes: Typhlopidae). 
Russian Journal of Herpetology, 6: 231-240. 

Lazell, J. D. 1999. The origins and evolution of the herpetofauna of the islands 
on the continental shelf of South China, pp. 79-96. In H. Ota (ed.). Tropical 
Island Herpetofauna: Origin. Current Diversity, and Conservation. Amster- 
dam. The Netherlands. Elsevier Science. 353 pp. 

Lazell, J. D., and W. Lu. 1990. Four remarkable reptiles from South China Sea 
islands. Hong Kong Territory. Asiatic Herpetological Research. 3: 64-66. 

Leviton, a. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards 
in herpetology and ichthyology: Part 1. Standard symbolic codes for institu- 
tional resource collections in herpetology and ichthyology. Copeia, 1985: 

LiM, K. K. P., and L. M. Chou. 1990. The heipetofauna of Singapore, pp. 49- 
59. In Essays in Zoology. Singapore. National University of Singapore. 

LiM. K. K. P., and F. L. K. Lim. 1992. A Guide to the Amphibians and Reptiles 
of Singapore. Singapore, Singapore Science Center. 160 pp. 

Macklin, S. 1988. Reclusive reptile has biologists bewildered. Hong Kong, South 
China Morning Post (28 May), 44(147): 1. 

McDiarmid. R. W.. J. A. Campbell, and T A. Toure. 1999. Snake Species of 


the World: A Taxonomic and Geographic Reference. Vol. 1. Washington, 
D.C., The Herpetologist's League. 511 pp. 

McDowell, S. B. 1974. A catalogue of the snakes of New Guinea and the Sol- 
omons, with special reference to those in the Bernice P. Bishop Museum. Part 
I. Scolecophidia. Journal of Herpetetology, 8: 1-57. 

OsHiMA, M. 1916. "A correction of the scientific names of Formosan snakes." 
Dobutsugaku Zasshi [= Zoological Magazine], 28: 84-86. 

Parker, H. W 1939. Reptiles and amphibians from Bougainville. Solomon Is- 
lands. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 15: 1-5. 

Peters, W C. H. 1874. Uber neue Reptilien (Peropus, Agama, Euprepes, Lygo- 
soma, Typhlops, Heterolepis) der herpetologischen Sammlung des Berliner 
zoologischen Museums. Monatsberichte der Koniglich Akademie der Wis- 
senschaften zu Berlin, 1874: 159-164. 

Peters, W C. H. 1881. Einige herpetologische Mittheilungen. 1. Uebersicht der 
zu den Familien der Typhlopes und Stenostomi gehorigen Gattungen oder 
Untergattungen. Sitzzungsberichte der Gesellschaft Naturforschenden Freun- 
de zu Berlin. 1881: 69-71. 

Romer, J. D. 1970. Revised annotated checklist with keys to the snakes of Hong 
Kong. Memoirs of the Hong Kong Natural History Society, 8: 1-22. 

Savage, J. M. 1950. Two new blind snakes (genus Typhlops) from the Philippine 
Islands. Proceedings of the California Zoological Club, 1: 49-54. 

Schlegel, H. 1839 in 1837-1844. Abbildungen Neuer Oder Unvollstandig Be- 
kannter Amphibien, Nach der Natur Oder dem Leben Entworfen, Herausge- 
geben und mit Einem Erlauternden Texte begleitet. Diisseldorf, Germany, 
Arnz & Comp. 80 pp. 

Smith, M. A. 1943. The Fauna of British India, Ceylon and Burma, Including the 
Whole of the Indo-Chinese Sub-region. Reptilia and Amphibia. Vol. Ill — 
Serpentes. London, Taylor and Francis. 583 pp. 

Sternfeld, R. 1913. Beitrage zur Schlangenfauna Neuguineas und der benach- 
barten Inselgruppen. Sitzungsberichte der Gesellschaft Naturforschender 
Freunde zu Berlin, 1913: 384-389. 

Stoliczka, F. 1871. Notes on some Indian and Bumese ophidians. Journal of the 
Asiatic Society of Bengal (Natural History), ser. 2, 40: 421-445. 

Taylor, E. H. 1918. Reptiles of the Sulu Archipelago. Philippine Journal of 
Science, 13D: 233-267. 

. 1947. Comments on Ceylonese snakes of the genus Typhlops with de- 
scriptions of new species. The University of Kansas Science Bulletin, pt. 2. 
31: 283-298. 

. 1962. New Oriental reptiles. The University of Kansas Science Bulletin, 

43: 209-263. 
. 1965. The serpents of Thailand and adjacent waters. The University of 

Kansas Science Bulletin, 45: 609-1096. 
Waite, E. R. 1897. A new blind snake from the Duke of York Island. Records 

of the Australian Museum, 3: 69-70. 
Waite, E. R. 1918. Description of a new blind snake from the Solomon Islands, 

20 BREVIORA No. 512 

with a list of species from the Austro-Malayan and Polynesian sub-regions. 

Records of the South Australian Museum, 1: 35-38. 
Wall, F. 1903. A prodromus of the snakes hitherto recorded from China. Japan, 

and the Loo Choo Islands; with some notes. Proceedings of the Zoological 

Society of London, 1903: 84-102. 
. 1909a. Notes on snakes from the neighbourhood of Darjeeling. Journal 

of the Bombay Natural History Society, 19: 337-357. 
. 1909b. A new blind snake from the western Himalayas. Journal of the 

Bombay Natural History Society, 19: 805-806. 
. 1913. Some new snakes from the Oriental region. Journal of the Bombay 

Natural History Society, 22: 514-516. 

Wallach, V. 1985. A cladistic analysis of the terrestrial Australian Elapidae, pp. 
223-253. In G. Grigg. R. Shine, and H. Ehmann (eds.). Biology of Austra- 
lasian Frogs and Lizards. Chipping Norton. Australia. Royal Zoological So- 
ciety of New South Wales. 527 pp. 

. 1993a. A new species of blind snake. Typhlops marxi, from the Philip- 
pines (Serpentes: Typhlopidae). Bulletin of the Raffles Museum, 41: 263- 

. 1993b. The supralabial imbrication pattern in the Typhlopoidea (Reptilia: 

Serpentes). Journal of Herpetology. 27: 214-218. 

. 1994. The status of the Indian endemic Typhlops aciitus (Dumeril and 

Bibron) and the identity of Typhlops psittacus Werner (Reptilia. Serpentes. 
Typhlopidae). Bulletin de ITnstitut Royal des Sciences Naturelles de Bel- 
gique. 64: 209-229. 

. 1995. A new genus for the Ramphotyphlops suhociilahs species group 

(Serpentes: Typhlopidae). with description of a new species. Asiatic Herpe- 
tological Research. 6: 132-150. 

. 1996. The systematic status of the Ramphotyphlops flaviventer (Peters) 

and related forms (Serpentes: Typhlopidae). Amphibia-Reptilia. 17: 341-359. 

. 1997. Two new blind snakes of the Typhlops ater species group from 

Papua New Guinea (Serpentes: Typhlopidae). Russian Journal of Herpetolo- 
gy. 3: 107-118. 

. 1998a. The lungs of snakes, pp. 93-295. In C. Gans and A. S. Gaunt 

(eds.). Biology of the Reptilia. Vol. 19 (Morphology G). Ithaca. New York. 
Society for the Study of Amphibians and Reptiles. 660 pp. 

. 1998b. The visceral anatomy of blindsnakes and wormsnakes and its 

systematic implications (Serpentes: Anomalepididae, Typhlopidae. Lepto- 
typhlopidae). Ph.D. Dissertation. Boston, Massachusetts. Northeastern Uni- 
versity. 61 1 pp. 

. 1999. Typhlops meszoelyi, a new species of blind snake from northeastern 

India (Serpentes: Typhlopidae). Herpetologica, 55: 185-191. 

. 2000. Critical review of some recent descriptions of Pakistani Typhlops 

by M. S. Khan. 1999 (Serpentes: Typhlopidae). Hamadryad. 25: 129-143. 

. 2001. Typhlops roxaneae, a new species of blind snake, with a synopsis 


of the Typhlopidae of Thailand (Serpentes: Scolecophidia). Raffles Bulletin 
of Zoology, 49: 39-49. 
. 2003. Scolecophidia miscellanea. Hamadryad, 27: 227-245. 

Wallach, v., and I. INEICH. 1996. Redescription of the rare Malagasy blind snake, 
Typhlops graudidieri Mocquard, with placement in a new genus (Serpentes: 
Typhlopidae). Journal of Herpetology, 30: 367-376. 

Werner, F. 1917. Uber einige neue Reptilien und einen neuen Frosch des Zool- 
ogischen Museums in Hamburg. Mitteilungen aus dem Zoologischen Muse- 
um in Hamburg (1916), 34: 31-36. 

Wynn, a. H., and a. E. Leviton. 1993. Two new species of blind snake, genus 
Typhlops (Reptilia: Squamata: Typhlopidae), from the Philippine Archipela- 
go. Proceedings of the Biological Society of Washington, 106: 34-45. 

Zhao, E.-M., and K. Adler. 1993. Herpetology of China. Oxford, Ohio, U.S.A., 
Society for the Study of Amphibians and Reptiles. 522 pp. 

Zhao, E.-M., M.-H. Huang, and Y. Zong. 1998. Fauna Sinica. Reptilia. Vol. 3. 
Squamata: Serpentes. Beijing, Science Press. 522 pp.