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ANNIVERSARY Of SCIENCES
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 22, pp. 381-392, 11 figs. November 14, 2003
The Genus Roboastra Bergh, 1877
(Nudibranchia: Polyceridae: Nembrothinae)
in the Atlantic Ocean
Marta Pola!, Juan Lucas Cervera!, and Terrence M. Gosliner
! Departamento de Biologia, Facultad de Ciencias del Mar y Ambientales, Universidad de Cadiz,
Apdo. 40, 11510 Puerto Real (Cadiz), Spain, marta.pola@uca.es; * California Academy of Sciences,
Golden Gate Park, San Francisco, CA 94118, USA, tgosliner@ calacademy.org.
The systematics of the phanerobranch dorid genus Roboastra Bergh, 1877 in the
Atlantic Ocean is reviewed. Roboastra europaea Garcia-Gomez, 1985 is redescribed
and its geographical range updated. Roboastra caboverdensis sp. nov. is described
from material from Cape Verde Archipelago. The two species differ in their color
pattern. The arrangement of the yellow lines or bands differs between the two
species with denser lines in R. europaea. The base of the rachidian radular tooth is
more curved in R. caboverdensis than in R. europaea and the upper cusp of the inner
lateral tooth is bifid in R. europaea while in R. caboverdensis it is simple.
Se revisa la sistematica del género del dérido fanerobranquio Roboastra Bergh, 1877
en el Océano Atlantico. Se redescribe Roboastra europaea Garcia-Gémez, 1985, asi
como se actualiza su distribuci6n geografica. Se describe Roboastra caboverdensis sp.
noy a partir de material procedente del archipiélago de Cabo Verde. Las dos especies
se diferencian por su patron cromatico. La disposicion de las lineas 0 bandas amar-
illas difiere entre las dos especies, con un mayor numero de ellas en R. europaea. La
base del diente radular raquideo esta mas curvada en R. caboverdensis y la cuspide
superior del diente lateral interno es bifida en R. europaea mientras que en R.
caboverdensis es simple.
The genus Roboastra was described by Bergh (1877). Until Burn’s revision (1967), this genus
included three species: Roboastra gracilis (Bergh, 1877) (type species), R. rubropapulosa (Bergh,
1905) and R. luteolineata (Baba, 1936), all with an Indo-Pacific distribution. Burn described a new
species, R. arika and suggested that R. rubropapulosa should be considered as a synonym of R.
gracilis. Some years later, Farmer (1978) described R. tigris from the eastern Pacific. Garctia-
Gomez (1985) then described R. europaea from the Strait of Gibraltar, the only species known
from Atlantic-Mediterranean waters. Thus, to date, the genus Roboastra includes five named
species. The only morphological data stem from the original descriptions, with the exception of the
redescription of R. gracilis by Burn (op. cit.) and its taxonomic comparison with R. luteolineata by
Hamatani and Baba (1976).
No additional studies have treated members of this genus except for that of R. tigris (Carté and
Faulkner 1983, 1986) and, more recently, in which R. europaea was the focus of molecular phylo-
genetic (Grande et al., 2002) and feeding ecology (Megina and Cervera 2003) studies. Recent col-
lections from the Cape Verde Archipelago (West Africa) have yielded several specimens of a sec-
ond undescribed Atlantic species of this genus.
381
382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 22
In this paper, we describe the new species from Cape Verde. We also redescribe R. europaea,
largely from material from the Iberian Peninsula, and update its geographical range.
MATERIAL AND METHODS
Specimens were dissected by dorsal incision. At least three specimens of each species were
examined anatomically. Their internal features were examined and drawn under a dissecting micro-
scope with a camera lucida. Particularly interesting soft parts were critical-point dried for scanning
electron microscopy (SEM). Special attention was paid to the morphology of the reproductive sys-
tem. The buccal mass was removed and dissolved in 10% sodiun hydroxide until the radula was
isolated from the surrounding tissue. The radula was then rinsed in water, dried, and mounted for
examination by scanning electron microscopy.
The materials examined are deposited in the California Academy of Sciences, San Francisco
(CASIZ), the Museo Nacional de Ciencias Naturales, Madrid (MNCN) and the Museu Municipal
de Funchal (Historia Natural) (MMEP).
SPECIES DESCRIPTIONS
Family Polyceridae Alder and Hancock, 1845
Subfamily Nembrothinae Burn, 1967
Genus Roboastra Bergh, 1877
Roboastra europaea Garcia-Gomez, 1985
(Figs. 1A, 2, 3A—-C, 4, 5, 6)
MATERIAL EXAMINED.— MNCN 15.05/46612. 1 specimen, 10 m depth, Torre, Marbella,
Spain, July 1995, J.L. Gonzalez, 30 mm. CASIZ 166049. 1 specimen, 10 m depth, Torre, Marbella,
Spain, September 1995, M.T.Barrea, 18 mm. CASIZ 166053. 2 specimen, Torre, Marbella, Spain,
August 1996, K.L. Schick, 10.7m, 15 y 19 mm. MNCN 15.05/29203, 1 specimen, La Herradura,
Granada, Spain, February 1993, A.Barrajon and M. Zarauz, 14 mm. MMF 31021, | specimen, 100
m depth, Funchal, Madeira, June 1999, 35 mm. MNCN 15.05/46613. 1 specimen, 20 m depth,
Ponta de Baleeira, Sagres, Portugal, July 2002, M.Pola, 14 mm. MNCN 15.05/46613. 1 specimen,
20 m depth, Ponta de Baleeira, Sagres, Portugal, July 2002, M.A. Malaquias, 10 mm. Specimens
were collected on rocks and were measured preserved.
DISTRIBUTION.— This species is known mainly from the Strait of Gibraltar and southern
Iberian Peninsula (Garcia-Gémez 1985; 2002!; Cervera et al. 1988; Garcia-Gémez et al. 1989,
1991; Moreno and Templado 1998; Schick 1998; Megina 2000; Ocana et al. 2000; Sanchez-Tocino
et al. 2000; Grande et al. 2002; Megina and Cervera 2003). One misidentified specimen of Plo-
camopherus trom Madeira deposited at the Natural History Museum (London) (Reg. No.
1863.9.19.3), supposedly collected by Rev. R. Lowe, was correctly identified as belonging to
Roboastra, very probably R. europaea. This conclusion has been strongly supported by the recent
collection of one specimen (also photographed) of this species at Funchal Harbour.
This species has been also recorded in southwestern Portugal (Calado et al. 2002) and
Catalunian coasts (northeastern Iberian Peninsula, Mediterranean Sea) (K.L. Schick, pers. com-
mun. ).
| The specimens described by Garcia~-Gémez (2002) are the same of those described by this author in 1985 for the orig-
inal description of this species.
POLA ET AL.: NUDIBRANCH GENUS ROBOASTRA IN THE ATLANTIC OCEAN 383
EXTERNAL MORPHOLOGY (Fig. 1A).— The body is elongate and limaciform. The preserved
animals are 10-40 mm in length. The body surface is lightly uneven with the edge of the mantle
not sharply angled. Foot is linear with a pointed posterior end of the foot. The head is rounded with
a pair of perfoliate rhinophores (bearing 30-35 lamellae) that are completely retractile into their
sheaths. The oral tentacles are well developed and grooved dorsolaterally along a part of their
A
a
Portugal. B. Roboastra caboverdensis sp. nov., specimen from Banco Joao Valente, Ilha da Boavista, Cape Verde.
384 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 22
length. There are five non-retractile, bipinnate gills surrounding the anal papilla and forming a half
circle; the three central gills are larger than the two lateral ones. The genital aperture is located mid-
way between the gills and rhinophores, on the right side. The body wall is highly muscular.
Regarding the color pattern, this species can exhibits two color phases. One of them has a grey or
bluish grey ground color; the other has dark blue ground color. Both phases have several yellow or
yellowish orange bands on the notum and both sides of the body. These bands are variable in num-
ber, shape, length and width. The rhinophores and oral tentacles are grey/bluish grey to dark blue.
The inner side of the gill rachis are yellow or yellowish orange. The outer side is frequently also
this color, but rarely may be a dark blue. The secondary pinnae are grey/bluish grey to dark blue.
The yellow/ yellowish orange areas are surroundend by a tiny violet line that is easily visible in
most animals, except in the darkest individuals. The rhinophoral sheaths are also grey/bluish grey
to dark blue (Fig. 1A).
INTERNAL MORPHOLOGY.— A general view of the internal anatomy can be seen in Figure 2.
The buccal mass is elongate and tubular, well developed with a pair of elongate pouches opening
into the digestive system at the junction of the oral tube and muscular pharynx (Fig. 3). The sali-
vary glands are short and thick, entering on the buccal mass and flanking the esophagus. The labi-
al cuticle lacks any armature. There is a well developed blood gland that is granular in texture. The
radular formula of two specimens of 15 mm length (preserved) is 23 x 4.1.1.1.4. and that of the 30
mm specimen (preserved) has the formula 25 x 4.1.1.1.4. (Fig. 4A—C). The rachidian tooth (Fig.
4B) is broad, thin and curved at its base with three well-differentiated cusps. The inner lateral tooth
nL gr ae
Figure 2 (left). Roboastra europaea Garcia-Gomez, 1985. General arrangement of the internal organs, au = auricle, bb
= buccal bulb, be = bursa copulatrix, bg] = blood gland, ca = cephalic artery, cg = cerebral ganglion, hg+dg = hermaphro-
dite gland+digestive gland, in = intestine, oe = oesophagus, ot = oral tube, pe = pericardium, m = rhinophoral nerves, vd =
vas deferens, ve = ventricle, vg = vestibular gland.
FIGURE 3 (right). Roboastra europaea Garcia~-Gémez, 1985. Detail of the oral tube and buccal mass.
POLA ET AL.: NUDIBRANCH GENUS ROBOASTRA IN THE ATLANTIC OCEAN 385
AVA
FiGuRE 4. Roboastra europaea Garcia-Gomez, 1985. CASIZ 166053, scanning electron micrographs of radula: A.
Half-row of radular teeth. B. Rachidian teeth. C. Inner lateral tooth.
(Fig. 4C) has a strongly curved bifid inner cusp. The inner branch of this cusp is thin and smaller
than the second. The outer cusp is undivided with a long spur-like denticle near the base. The
remaining lateral radular teeth are quadrangular and lack cusps or denticulation and become small-
er near the margin.
REPRODUCTIVE SYSTEM (Fig. 5).— The hermaphroditic duct widens into a S-shaped ampulla
which has thick walls. The bursa copulatrix is rounded and larger than the seminal receptacle,
which is elongate. The seminal receptacle has a short duct that connects to the vagina near the
bursa. The deferent duct, which lacks a morphologically well-differentiated prostate, is long and
coiled and ends in a dilated penial atrium. The vestibular gland is large with muscular walls, con-
vex on one side and concave on the other. The penis is located within the distal end of this muscu-
lar portion, and it is armed with, at least, three different kinds of hooked and chitinous spines
arranged in helicoidal rows. The types of spines and their arrangement on the penis are shown in
Figure 6.
386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 22
FIGURE 5. Roboastra europaea Garcia-Gomez, 1985. Reproductive system, am = ampulla, be = bursa copulatrix, fgm
= female gland mass, hd = hermaphrodite duct, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = vas
deferens, vg = vestibular gland.
FIGURE 6. Roboastra europaea Garcia-Gomez, 1985.
CASIZ 166053, scanning electron micrographs of penis: A.
Protruded penis and penial armature. B. Detail of distal
spines. C. Detail of middle spines.
POLA ET AL.: NUDIBRANCH GENUS ROBOASTRA IN THE ATLANTIC OCEAN 387
Roboastra caboverdensis Pola, Cervera, and Gosliner, sp.nov.
(Figs. 1B, 7, 8, 9A-C, 10, 11A—D)
TYPE MATERIAL.— HOLOTYPE: CASIZ 166047, 1 specimen, 38 m depth, 15 miles NW Santo
Antao Island, Cape Verde Archipelago, July 2002. PARATYPES: MMF35083, 1 specimen, Tarrafal
Island. Cape Verde Archipelago. December 1998, P. Wirtz, 25 mm. MNCN. 15.05/46614, 2 speci-
men, 20 m depth, Banco Joao Valente, Boavista Island, Cape Verde Archipelago, August 2002,
M.A. Malaquias, 26/30 mm (70 mm in life). CASIZ 166052, 2 specimen, 30 m depth, Banco Joao
Valente, Boavista Island, Cape Verde, August 2002, M.A. Malaquias, 31/37 mm (70 mm in life).
MNCN. 15.05/46617. 3 specimens, Boavista Island, August 2002, C. Grande, 13/15/18 mm.
CASIZ 166050, 1 specimen, Sao Vicente, October 2002, G. Calado, 20mm. Specimens were col-
lected on rocks and were measured preserved.
ETyMOLoGy.— The name caboverdensis refers to the Cape Verde Archipelago where this
species is found.
DISTRIBUTION.— Thus far, known only from Cape Verde Archipelago.
EXTERNAL MORPHOLOGY.— The body is elongate and limaciform with a long and pointed
posterior end of the foot. The preserved animals are 10-40 mm in length. The living animals (Fig.
1B) may reach 70 mm in length. The body surface is strongly wrinkled. The foot is linear. The head
is rounded with a pair of conical, completely retractile, perfoliate rhinophores with approximaly 35
tightly packed lamellae. The oral tentacles are strongly developed and dorsolaterally grooved along
a part of their length. There are five non-retractile tripinnate gills, with the three anteriormost being
more highly developed. The gills form a semicircle surrounding the anal papilla. The genital pore
opens on the right side, midway between the gills and rhinophores. The ground color is dark blue,
almost black. A wide yellow submarginal band follows the inner notal edge. This band is interrupt-
ed in some specimens. A second yellow band arises from the former, just in front of both
rhinophores and surrounds the inner side of their sheaths, continuing to the rear to the base of the
gills. These bands can bifurcate at their origin. In this case, the sheath of the rhinophores is sur-
rounded by the shorter branch. In either situation, these bands surround the gill to join each other
posteriorly. These last bands can be interrupted or continuous. The edge of foot is also bordered by
a wide yellow band. On the sides of the body, just below the notal edge, there is a line of the same
colour that bifurcates and surrounds the genital pore, continuing to the end of the posterior end of
the foot. Both branches can be continuous or not. Moreover, several yellow lines, varying in num-
ber and length, are arranged between the dorsal and the foot bands. The oral tentacles, the
rhinophores and the posterior part of their sheath are also blue-black. The gills are blue-black, but
the inner and outer sides of the rachis of each one have a yellow line between them.
INTERNAL MORPHOLOGY.— The general arrangement of the internal organs is shown in Figure
7. The anterior digestive tract begins with a long thick-walled muscular oral tube, that continues
into the buccal mass. At their junction, a pair of elongate pouches open into the digestive system
(Fig. 8). There is a pair of small, short and wide salivary glands on the buccal mass, flanking the
esophagus. The radular formula of two specimens of 70 mm (in life) is 33 x 3-4.1.1.1.3-4 (Figs.
9A-C). The rachidian tooth (Fig. 9B) is broad, clearly curved at the base, having three well-differ-
entiated cusps. The inner lateral tooth (Fig. 9C) is hooked with two well developed elongate cusps.
The inner one is very long, having sharp and curved edges on the internal side and a prominent pro-
jection (see the arrows Fig. 9A) on its outer edge. The outer lateral teeth (3 to 4) are smaller and
quadrangular without prongs, and decreasing in size from the inner to the outer side of the radula.
A labial cuticle is present, but lacks armature.
REPRODUCTIVE SYSTEM.— The reproductive system is shown in Figure 10. The hermaphro-
388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 22
ve
hg+dg ____\
Figure 8. Roboastra caboverdensis sp. nov. Details of
the oral tube and buccal mass, bb = buccal bulb, bg = buc-
cal ganglion, oe = oesophagus, ot = oral tube.
FIGURE 7. Roboastra caboverdensis sp. noy. General
arrangement of the internal organs. au = auricle, bb = buccal
bulb, bgl = blood gland, ca = cephalic artery, cg = cerebral ditic duct has an “S” shaped ampulla that con-
ganglion, fm = female gland, hg+dg = hermaphrodite tjnues into the spermoviduct. The vas deferens
glad dieestive late, ua = intestine, oe = Cesophaeus of = is lone and coiled. with a unitGmneeidtimlene
oral tube, pe = pericardium, rn = rhinophoral nerves, ve = = L
ventricle: slightly narrower in the prostatic part. It ends in
a dilated penial section. The penis is armed
with, at least, three different kind of spines arranged in helicoidal rows. Types of spines and their
arrangement on the penis are shown in Figures 11A—D. The bursa copulatrix is rounded and the
seminal receptacle elongate. The seminal receptacle joins with the vagina, near the bursa via a short
duct. The vagina is short and straight, opening into the genital atrium, near the vestibular gland.
This gland is very well developed, flattened, with muscular walls.
DISCUSSION
In 1985, Garcia-Gomez described the first species of the genus Roboastra in the Atlantic
Ocean, from the Strait of Gibraltar. The original description of the external anatomy, radula and
reproductive system of Roboastra europaea is in agreement with our specimens of this species;
however, our specimens exhibit a different pattern of coloration than has been previously
described for this species (Schick 1998; Megina 2000; Garcia~-Gémez 2002:224, phot. 447; Sanchez
2001). Moreover, it is confirmed that the spicules of the surface of the mantle mentioned in the
original description do not exist (perhaps they were an artifact of preservation). A pair of small,
short and wide salivary glands on the buccal mass flanking the esophagus are described for the first
time. These salivary glands are present in the other two genera of the subfamily Nembrothinae,
? The colour pattern supplied by Garcfa~-Gémez (2002) is an adaptation from his original description (Garcia-Gémez,
1985), with additional and later information, although the specimens included in the “Material” section are the same in both
references.
POLA ET AL.: NUDIBRANCH GENUS ROBOASTRA IN THE ATLANTIC OCEAN
FIGURE 9. Roboastra caboverdensis sp. nov. CASIZ 166052, scanning electron micrographs of radula. A. Half-row of
radular teeth. B. Rachidian teeth. C. Inner lateral tooth.
Nembrotha and Tambja, but in these they are longer and more robust than in Roboastra. Garcia-
G6émez (1985) did not describe the presence of a pair of elongate pouches that open into the diges-
tive system at the junction of the oral tube and muscular pharynx. The function of these structures
is still unknown; nevertheless, Burn (1967) described similar structures in Roboastra gracilis.
The external and internal features of Roboastra caboverdensis permit us to distinguish it from
its congeneric Atlantic species. The arrangement of the yellow lines or bands in both species is dif-
ferent and are more numerous and tightly packed in R. europaea. Moreover, R. europaea has two
colour phases, light and dark (see Cervera et al. 1988 and Ocajia et al. 2000, for a colour picture of
the light phase).
Regarding the internal anatomy of both species, the base of the rachidian radular tooth is more
390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
curved in R. caboverdensis and the upper cusp of
the inner lateral radular tooth is bifid in R.
europaea while in R. caboverdensis it is simple.
The reproductive system of R. caboverdensis is
similar to that described for R. europaea except that
the portion of the ejaculatory duct near the prostate
is more highly convoluted in R. europaea. The
penis in both cases 1s armed and the spines, of three
different sizes and shapes, are implanted in heli-
coidal rows. In both R. europaea and R. caboverd-
ensis, the distal two-thirds of the penis has elon-
gate, curved spines, and the basal spines are medi-
um sized but straighter than the distal ones. Be-
tween them, there are some rows of short, curved
spines, a few with little spines just behind them.
There is another dark blue/black with yellow
bands or lines nembrothid in the Cape Verde Archi-
pelago, Tambja simplex Ortea and Moro, 1998.
However, the yellow pattern of this species has few
lines with a different arrangement, and the internal
anatomy (salivary glands, labial cuticle, radular
Volume 54, No. 22
pr
am
FIGURE 10. Roboastra caboverdensis sp. nov. Re-
productive system, am = ampulla, be = bursa copulatrix,
fgm = female gland mass, hd = hermaphrodite duct, p =
penis, pr = prostate, rs = receptaculum seminis, v =
vagina, vd = vas deferens, vg = vestibular gland.
teeth and reproductive system) are characteristic of Zambja (Ortea and Moro 1998; Cervera et al.
2000).
10um
pall
C. Detail of middle spines. D. Detail of basal spines.
FiGURE 11. Roboastra caboverdensis sp. nov. MNCN 15.05/46614, A. Protruded penis. B. Detail of distal spines.
10um
==
POLA ET AL.: NUDIBRANCH GENUS ROBOASTRA IN THE ATLANTIC OCEAN 3)
ACKNOWLEDGMENTS
Our most sincere gratitude to Drs. Gongalo Calado, Peter Wirtz, Manuel A. Malaquias and
Cristina Grande, for providing us the Cape Verde specimens and their photographs, to Dr. César
Megina, Karl L. Schick and Antonio D. Abreu (MMP) for giving us kindly their unpublished data
and photographs of Roboastra europaea and R. caboverdensis. To Dr. David Reid (NHM) and
Antonio D. Abreu for sending the material from Madeira. We also express our gratitude to Mr. José
Maria Geraldia and Mr. Juan Gonzalez (from the Electron Microscopy Service of the University of
Cadiz) and Mr. Scott Serata (from the Electron Microscopy Laboratory of the California Academy
of Sciences) for providing facilities to take Scanning Electron Microscope photographs. Marta Pola
(M.P.) deeply thanks Yolanda Camacho-Garcia her assistance during the stay at the California
Academy of Sciences.
Finally, this paper has been benefited by the following projects REN2001-1956-C17-02/GLO
(Spanish Ministry of Science and Technology), REN2000-0890/GLO (Spanish Ministry of Science
and Technology) and PEET Grant DEB-9978155 (National Science Foundation, USA). One of the
authors (M.P.) has been benefited by a predoctoral fellowship and a grant for a short stay at the
California Academy of Sciences, both funded by the Spanish Ministry of Education, Culture and
Sports.
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GARCIA-GOMEZ, J.C., J.L. CERVERA, F.J. Garcia, J.A. ORTEA, S.F. GARCIA, A. MEDINA, AND L.P. BURNAY.
1991. Resultados de la Campania Internacional de Biologia Marina “Algarve 88”: Moluscos Opistobran-
quios. Bollettino Malacologico 27:125—138.
GRANDE, C., J. TEMPLADO, J.L. CERVERA, AND R. ZARDOYA. 2002. The complete mitochondrial genome of the
nudibranch Roboastra europaea (Mollusca: Gastropoda) supports the monophyly of opistobranchs.
Molecular Biology and Evolution 19 (10):1672-1685.
HAMATANI, I., AND K. BABA. 1976. Taxonomical comparison between the nudibranch species Roboastra gra-
cilis and R. luteolineata from Yoron Islands of the Amami Islands, Southren Japan. Venus 35 (3):135—137.
Meaina, C. 2000. Dieta y Especializacion Trofica en Moluscos Rudibranquios. Ph.D. Thesis. Universidad de
Cadiz. 157 pp.
MEGINA, C. AND J.L. CERVERA. 2003. Diet, prey selection and cannibalism in the hunter opisthobranch
Roboastra europaea. Journal of the Marine Biological Association of the United Kingdom 83:485—495.
Moreno, D. AND J. TEMPLADO. 1998. Nuevas aportaciones al conocimiento de los opistobranquios del sureste
espanol. I. Iberus 16(2):39-58.
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Verde. Avicennia 8/9:149-154.
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_~
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23, pp. 393-406, 12 figs. November 14, 2003
Redescription of Halgerda graphica Basedow and Hedley, 1905,
with Observations on External Morphological Variation within
Selected Species of Halgerda (Mollusca: Nudibranchia)
Shireen J. Fahey and Terrence M. Gosliner
Department of Invertebrate Zoology and Geology
California Academy of Sciences
Golden Gate Park, San Francisco, California 94118, USA
Halgerda graphica Basedow and Hedley, 1905 is redescribed based on examination
of two specimens; one collected in 1904 from Middle Harbour, Sydney, Australia,
housed at the Australian Museum, Sydney, and another specimen collected in 1990,
from Port Moorowie, near the type locality (Kangaroo Island, South Australia). The
reproductive system is described and illustrated for the first time. This species has
frequently been misidentified due to having similar external morphology to other
Halgerda species. A comparison is made to those species. Halgerda graphica has a
unique combination of external and internal characters that confirm it as a distinct
Halgerda species. The external characters include a “hieroglyphic” pattern of yellow
and black markings on the notum, small, similar-size dark spots on the ventral sur-
face, a small, sparse, dark-colored gill and rhinophores with a white base, dark tip
and a dark line on the posterior side. The external color variations of Halgerda
dichromis Fahey and Gosliner, 1999, H. okinawa Carlson and Hoff, 2000 and H. wil-
leyi Eliot, 1904 are also described, illustrated and compared to externally similar
species. It is the unique combination of external morphological characters such as
the color and pattern on the notum, the structure and color of the gills and
rhinophores that help to distinguish each species, although examination of internal
morphology can confirm the identification.
Basedow and Hedley (1905) described the nudibranch Halgerda graphica from two specimens
dredged off Antechamber Bay, Kangaroo Island, South Australia. They had at the time examined a
third specimen, which, although not included in the original description, they indicated that it
belonged to their new species. The original description of the external and radular morphologies of
the specimens were quite detailed, but the authors did not provide a description of the reproductive
morphology. This species did not show up again until 1990. In the meantime, other authors (e.g.,
Coleman 1975, 2001; Kay 1979; Kay and Young 1969; Wells and Bryce 1993) have erroneously
attributed the name Halgerda graphica to other species.
The present study describes two additional specimens of Halgerda graphica; the single spec-
imen mentioned by Basedow and Hedley, collected in approximately 1904 at Middle Harbour,
Sydney, and an additional specimen collected by N. Holmes in 1990 from Port Moorowie, Yorke
Peninsula, South Australia.
The nudibranch genus Halgerda Bergh, 1880 has been studied extensively in recent years
(Rudman 1978; Willan and Brodie 1989; Carlson and Hoff 1993, 2000; Gosliner and Fahey 1998;
! Contact author: sfahey @ calacademy.org
393
394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
Fahey and Gosliner 1999a, 1999b, 2000, 2001a, 2001b). Since 1998, the number of described
species increased from 14 to 35. Among Halgerda species are several that display external color
variation in specimens from similar habitats and from varying geographic ranges. Four species, in
particular, seem to be most commonly confused: Halgerda graphica, H. willeyi, H. okinawa
Carlson and Hoff, 2000, and H. dichromis Fahey and Gosliner, 1999. The present study illustrates
the color variations that can cause misidentification of these four species in the field and then
makes comparisons with the externally most similar species. Field notes and photographic records
of many observers, both amateur and professional, provide the basis of the discussion on external
color variation presented in this paper.
DESCRIPTIONS OF TAXA
Labiostomata Valdés, 2002
Family Discodorididae Bergh, 1891
Genus Halgerda Basedow and Hedley, 1905
(Type species: Halgerda formosa Bergh, 1880, by monotypy)
Halgerda graphica Basedow and Hedley, 1905
(Figs 1-3)
TYPE MATERIAL.— The type material is probably lost; it is not at the Australian Museum. The
type locality is Kangaroo Island, South Australia. The authors (Basedow and Hedley 1905) men-
tion an additional specimen deposited at the Australian Museum, Sydney (C18168, Location
#016276, collector not named) from Middle Harbour near Sydney, in September 1904. This spec-
imen was examined and is both described below and hereby designated as the neotype.
OTHER MATERIAL EXAMINED.— South Australian Museum, TD16542, one specimen, dissect-
ed, Port Moorowie, Yorke Peninsula, South Australia. 10 m deep, collected by Nigel Holmes, 15
December 1990.
EXTERNAL MORPHOLOGY.— Because there are no photographs or drawings available for spec-
imen C18168 (1904), the following is a description of the specimen collected in 1990 by N.
Holmes. This specimen is nearly identical to the drawing and description of Halgerda graphica
provided by Basedow and Hedley in 1905 (Fig. 1A).
The preserved animal is 28 mm in length. The body profile is rounded, convex (Fig. 1B) and
the dorsum has a low-ridged pattern. There are no small marginal tubercles. There is a low central
ridge running the length of the dorsum that splits into several smaller ridges as it nears the gill
pocket. The ridges have orange-yellow crests. The background color of the dorsum is gray-white.
Between the ridges are black spots with some spots circled or semi-circled with the same yellow
color as the ridges. The dark spots closest to the mantle edge are smaller than those on the dorsum.
The mantle margin is translucent white when viewed dorsally, but when viewed ventrally, a yellow
margin is apparent. On the underside of the mantle and along the side of the foot are dark spots of
various sizes. The foot margin is yellow (Fig. 1C). The oral tentacles are long and tapered.
The long rhinophores have a bulging club that is tapered at the tips. The club is angled poste-
riorly and there is dark brown to black coloration around the top half of the club up to the tip. The
base is translucent white and there is a dark line on the posterior side of the rhinophores that
extends from the base to the tip.
The bipinnate gill lies flat over the dorsum and is moderately pinnate. Each of the four main
gill rachae has a brown stripe on the anterior sides. The anal papilla is long and is the same color
as the body.
FAHEY AND GOSLINER: HALGERDA GRAPHICA 395
Vol XXDC Plate 10
Desedow del of pina. a¢ ast
HALGERDA GRAPHICA, Basedow & Hedley.
FicurE 1. A. Halgerda graphica Basedow and Hedley,
1905, from the original color plate. B. Halgerda graphica
South Australian Museum, (TD16542), photographed and
collected by N. Holmes, 1990. Dorsal view. C. Ventral
view.
BUCCAL ARMATURE.— The buccal mass
has dark spots. The labial cuticle is smooth
and devoid of any jaw rodlets. The radular
sac is elongate and extends well behind the
posterior end of the buccal mass. The radular
formula of the specimen collected in 1990 is:
43x30.0.30 (TD16542) (Fig. 2A). The radu-
lar formula of the specimen collected in 1904
could not be determined due to poor preser-
vation and deterioration of the radula. The
three outer teeth are much smaller than the
inner and middle lateral teeth and the outer
two teeth have tiny denticles (Fig. 2B). The 8
or so inner lateral teeth are smaller and have
shorter hooks than the middle lateral teeth
(Fig. 2C) and are arranged in a shallow V-
shaped pattern in the center of the radula.
The middle lateral teeth are hamate (Fig. 2D)
with long, pointed hooks. They have a flat-
tened flange, which overlaps the adjacent
tooth.
REPRODUCTIVE SYSTEM.— The repro-
ductive system is triaulic (Fig. 3). The long
ampulla is tubular, curved into a complete
loop and protrudes away from the bursa and
prostate. The ampulla narrows into the post-
ampullary duct, which bifurcates into the vas
deferens and oviduct. The long oviduct
enters the female gland mass. The female
gland mass is about the same size as the
bursa copulatrix. The long vas deferens sep-
arates from the ampulla and widens into the
glandular prostate. The prostate consists of
two distinct glandular types and they are well
differentiated as in most other members of
Halgerda. The muscular portion of the defer-
ent duct leaves the distal prostate in a long
duct that curves into one loop and multiple
half-loops, then enters the wide penial bulb.
The long uterine duct emerges from the
female gland mass and joins the ovoid recep-
taculum seminis near its base. The duct con-
396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
—)
Wows
# ig
FIGURE 2. Radular morphology of Halgerda graphica (TD16542). A. Entire radula. Scale = 2 um. B. Outer lateral teeth.
Scale = 50 um. C. Inner lateral teeth. Scale = 100 um. D. Middle lateral teeth. Scale = 20 um.
necting the receptaculum and the bursa is long and coiled. The receptaculum seminis is much
smaller than the thin-walled spherical bursa copulatrix. It lies under the bursa, but is not embedded
in the prostate. The prostate does not completely cover the bursa copulatrix as is common in other,
more highly derived species of Halgerda, but lies in a thin layer over two-thirds of the bursa. The
vaginal duct that emerges from the base of the bursa copulatrix is long and thin. Near its exit into
the bulbous vagina that is adjacent to the base of the penial sheath, is a muscular sphincter. The
vagina has long folds in the walls and tubercular glands on the exterior. The common genital aper-
ture is wide, large and has long fleshy folds that extend through the body wall. The opening of the
female gland mass is adjacent to the genital aperture.
—
FAHEY AND GOSLINER: HALGERDA GRAPHICA 397
DISCUSSION
Although Basedow and Hedley (1905)
described the external and radular morphology
of Halgerda graphica, these authors did not
illustrate or describe the reproductive morphol-
ogy. Despite this, based on the unique color pat-
tern of this species, there is little doubt that the
specimens examined for this study are
Halgerda graphica. Other authors (Coleman
1975, 2001; Kay 1979; Kay and Young 1969;
Wells and Bryce 1993) have erroneously attrib-
uted the name Halgerda graphica to other
species. Wells and Bryce, mislabeled a speci-
men of Halgerda gunnessi as H. graphica
Basedow and Hedley, 1905; Kay misidentified
a specimen of H. terramtuentis as H. graphica;
__ Ficure 3. Reproductive morphology of Halgerda graph- and Coleman (1975, 2001) misidentified a
See ee eee serena & specimen of H. willy as H. graphic
um, p = penis, rs = receptaculum seminis, v = vagina. Scale Examination of the reproductive morphol-
= 0.25 mm. ogy reveals similar characters to other
Halgerda such as a two-part prostate, a long,
convoluted deferent duct, a bulbous penial sheath and a wide, muscular vagina. Radular characters
also have similarities to other Halgerda species. Those characters are hooked mid-lateral teeth,
finely denticulate outer teeth that are much smaller than the remaining teeth, and small inner later-
al teeth.
Because of the similarity in external coloration of Halgerda graphica to H. gunnessi Fahey
and Gosliner, 2001, H. johnsonorum Carlson and Hoff, 2000 and H. willeyi Eliot, 1904, these three
are herein compared and contrasted to H. graphica. However, because Carlson and Hoff (2000)
have already compared H. graphica to H. okinawa, we will not repeat what they have already done.
Neither will we repeat what Fahey and Gosliner said when they compared H. gunnessi to H. john-
sonorum and to H. formosa Bergh, 1880, nor what Carlson and Hoff (2000) had to say when they
compared H. johnsonorum to H. willeyi. Rather, we encourage reference to their respective papers.
Here we concentrate on comparing the external morphology of Halgerda graphica to its most sim-
ilar species.
All four species of greatest concern to us, Halgerda graphica, H. gunnessi, H. johnsonorum
and H. willeyi, have a white or gray-white ground color with yellow to yellow-orange ridge crests.
Halgerda graphica has low ridges without tubercles as do H. gunnessi and H. johnsonorum. Only
H. willeyi has prominent tubercles. Although both H. graphica and H. johnsonorum have dark
spots or markings in the ridge concavities, only H. graphica has the “hieroglyphic markings”
described by Basedow and Hedley. These markings consist of a dark spot at the center of the con-
cavity surrounded by dark circles and lines (Fig. 1). The other two species, H. gunnessi and H. wil-
leyi may also have dark lines or markings, but both lack the associated spots.
The coloration of the mantle edge also distinguishes these four species. Halgerda graphica
does not have perpendicular dark markings along the mantle edge, but the edge markings of H.
johnsonorum appear as continuous lines, which extend upward and into the ridge concavities on
the dorsum. The dark perpendicular lines on the mantle margin of H. willeyi also extend up into the
398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
ridge cavities, but they are more numerous when compared to H. graphica. Halgerda gunnessi
does not have marginal lines (see Fahey and Gosliner 2001).
The ventral surfaces of the four species are not similar in that there are irregularly scattered
dark spots without lines only on Halgerda graphica. Halgerda johnsonorum and Halgerda willeyi
have black lines; H. gunnessi has no markings on its ventrum.
The coloration on the rhinophores and gills differs among the species. Halgerda graphica has
dark coloration on the tips of the rhinophores and a dark line on the posterior side. This is similar
to H. gunnessi. The other species have either dark spots (H. johnsonorum, H. willeyi) or dark spots
plus a dark stripe on the rhinophores (H. johnsonorum) and large gills. The gill of H. graphica is
small and darkly colored. The gill of H. willeyi is sparsely pinnate with dark speckles. The gill of
H. johnsonorum is large and has dark spots, and the gill of H. gunnessi is large, feathery and has
dark lined branches and a dark tip.
With regard to the internal morphology, Halgerda graphica has radular characters similar to
those of H. gunnessi and H. willeyi. All three also have three small outer teeth. H. johnsonorum is
distinct, having six outer teeth, with the penultimate being bifid. Halgerda graphica has two fine-
ly denticulate outer lateral teeth; the outer three of H. willeyi and H. gunnessi are not denticulate.
The reproductive morphology of Halgerda graphica is most similar to H. willeyi. The obvious
differences between the two are that the deferent duct of H. graphica is much longer and more con-
voluted than in H. willeyi and the vaginal duct of H. graphica is much wider with the vagina, being
much larger with tubercular glands on the exterior. A glandular vagina is not found in any of the
other three species. Both Halgerda graphica and H. gunnessi have a vaginal sphincter.
The present study confirms the combination of external and internal morphological characters
that identify Halgerda graphica and distinguish it from the four externally most similar species.
The external characters that distinguish this species in the field are: the “hieroglyphic” yellow and
black markings on the dorsum, no dark lines on the mantle edge, dark-tipped rhinophores with a
posterior medial line, a small, dark gill and small, dark spots on the ventral surface.
Halgerda dichromis Fahey and Gosliner, 1999
(Figs. 4-5)
MATERIAL EXAMINED.— V8234, one specimen, dissected, 42 mm, Scottburgh, Kwazulu,
Natal, South Africa, 25 m deep, collected by V. Fraser, 15 January 2000; V8233, one specimen, dis-
sected, 20 mm, Park Rynie, KwaZulu, Natal, South Africa, 25 m deep, collected by V. Fraser, 28
December 1999; V8232, one specimen, dissected, 16 mm, Park Rynie, KwaZulu, Natal, South
Africa, 25 m deep, collected by V. Fraser, 21 January 2000.
EXTERNAL MORPHOLOGY.— The external morphology of the specimens examined for this
study are as described by Fahey and Gosliner with some color variation. The variation includes the
presence of dark half-lines or spots on the dorsum of some specimens, in place of a heavy, dark
line. The more juvenile specimens may not have any dark markings at all. Variations in the exter-
nal color within this species are shown in Figures 4A-C.
RADULAR MORPHOLOGY.— There were no differences noted in the radular morphology
between the recent specimens examined (Figs SA—D) and Fahey and Gosliner’s (1999) original
description and line drawings. Figure 5A—D are the first SEMs of the radula of this species.
REPRODUCTIVE SYSTEM.— There were no differences in the sexually mature specimens exam-
ined for this study (Fig. 6) and Fahey and Gosliner’s (1999) original description of Halgerda
dichromis.
REMARKS.— Halgerda dichromis was described from a single specimen collected in 1980
FAHEY AND GOSLINER: HALGERDA GRAPHICA 399
«<* >
“1 make [ad
FiGure 4. Color variation of Halgerda dichromis. A-C, Photographed and collected by
V. Fraser, 2000.
from Durban Harbor, South Africa. Since the original description, additional specimens have been
collected and/or photographed from the same locality (present study) and, thus, allow a further
examination of the species.
Halgerda dichromis has a variable external coloration (Figs. 4A—C). The coloration of the
holotype includes orange and black lines that form a reticulate pattern on the notum (Fahey and
Gosliner 1999). Other patterns include having only orange or yellow lines with dark lines or
splotches and without dark markings at all, particularly on more juvenile specimens.
Halgerda dichromis externally is most similar to H. formosa (see Fahey and Gosliner 1999 for
details).
Halgerda okinawa Carlson and Hoff, 2000
(Figs. 6-8)
MATERIAL EXAMINED.— CASIZ 144092, one specimen, 80 mm, dissected. Izu Peninsula,
Japan. 22 m depth, collected by R. Nakano, April 2000; CASIZ 144093, one specimen, 46 mm, dis-
sected. Izu Peninsula, Japan. 20 m depth, collected by R. Nakano, May 2000; CASIZ 144097, one
specimen, 50 mm, dissected. Izu Peninsula, Japan. 22 m depth, collected by R. Nakano, April 2000.
EXTERNAL MORPHOLOGY.— The external morphology of the specimens examined from the
Izu Peninsula have the same range of variation as noted in the original description of H. okinawa
(Carlson and Hoff, 2000). Those variations include number, length and width of the dark streaks,
number of lines on the inner surface of the branchia and presence of a yellow mantle margin on the
specimens examined for this study. One of the specimens we examined from the Izu Peninsula has
a paler shade of white on the dorsum with pale yellow tubercles (Fig. 7A). The other specimen (Fig.
400
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
FIGURE 5. Radular morphology of Halgerda dichromis (V8233). A. Inner lateral teeth. Scale = 10um. B. Middle lateral
teeth. Scale = 50.8 um. C. Middle lateral teeth. Scale = 50.8 um. D. Outer lateral teeth. Scale = 10 um.
FIGURE 6. Reproductive morphology of Halgerda
dichromis (V8234). Abbreviations: am = ampulla, be =
bursa copulatrix, dd = deferent duct, fgm = female gland
mass, ga = genital atrium, p = penis, rs = receptaculum sem-
inis, V = vagina. Scale = 0.8 mm.
7B) matched the original description from
Okinawa. A specimen from Lembeh Strait (Fig.
7C) had few dark lines but more spots that the
more commonly found specimen (Figs 7D-E).
RADULAR MORPHOLOGY.— There were no
differences noted in the radular morphology
between the recent specimens examined (Fig.
8) and Carlson and Hoff’s (2000) original
description.
REPRODUCTIVE | MORPHOLOGY.— The
reproductive morphology of the specimens
examined for the present study (Fig. 9) were
nearly identical to the original description . The
exception is that the vagina is wider in the
specimens we examined than was illustrated
and drawn by Carlson and Hoff (2000).
REMARKS.— Since the original descrip-
tion of H. okinawa, additional specimens col-
lected from the Izu Peninsula, Japan and pho-
FAHEY AND GOSLINER: HALGERDA GRAPHICA 401
(C) $. KATO
FIGURE 7. Color variation of Halgerda okinawa. A. Photographed by Hachijo. B. Photographed by S. Kato, 2001.
C. Photographed by Carine Scheurs. D. Photographed by Carlson and Hoff. E. Photographed by R. Bolland.
tographed elsewhere show variation in the external color pattern (Fig. 7C—D). Differences between
Halgerda okinawa and H. graphica were discussed thoroughly by Carlson and Hoff (2000).
402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
Lin 200pn1 EXT © 15.00k% WO* iin Fle blame = HaBehole-14.0F
| eae.
= tmm File Narre = MeBeuter-12 of
NI ges
om
EXT e 1540kf WOe Sm Fe Name « MaDe
vA
i & Dp. my Y, : (gry
FIGURE 8. Rad of the radula. Scale = 200 um. B. Outer lat-
eral teeth. Scale = 10 um. C. Inner lateral teeth. Scale = 20 um. D. Middle lateral teeth. Scale = 30 um.
Halgerda willeyi Eliot, 1904
(Figs 9-11)
= Halgerda willeyi in Coleman, 2001, p. 58, center photo, AMPI 117; and in Coleman, 1975 p. 63, Plate 170.
MATERIAL EXAMINED.— CASIZ 144095, two specimens: 35 mm, 51 mm (dissected). 20 m
depth, collected by R. Nakano, March 2000; CASIZ 144096, three specimens; 25mm, 28 mm, 31
mm (dissected). 21 m depth, collected by R. Nakano, April 2000; CASIZ 144121, one specimen,
40 mm, dissected. 41.5 m depth. 1.3 km ENE Maeki-zaki, Seragaki, Okinawa, Ryukyu Islands,
Japan, collected by R. Bolland, January 2001; CASIZ 144123, one specimen, 30 mm. 41.5 m
depth. 1.3 km ENE Maeki-zaki, Seragaki, Okinawa, Ryukyu Islands, Japan, collected by R.
Bolland, 8 December 2000; CASIZ 134919, one specimen, 74 mm. 43 m depth. 1.3 km ENE
Maeki-zaki, Seragaki, Okinawa, Ryukyu Islands, Japan, collected by R. Bolland, 17 August 2000;
BMNH, one specimen, 60 mm. Intertidal. Bapper Bay, Aden, Yemen, leg. Sgt. Howse, RAMC.
October 1966.
EXTERNAL MORPHOLOGY.— Both specimens examined for this study are externally similar to
other specimens of H. willeyi published elsewhere. However, this species displays great variation
in external coloration (Figs. 1OA—D; also see Gosliner et al. 1996; Marshall and Willan 1999; Ono
1999: Coleman 1975, 2001; Bolland 2003; Rudman 2003).
RADULAR MORPHOLOGY.— There were no differences noted in the radular morphology
FAHEY AND GOSLINER: HALGERDA GRAPHICA 403
FIGURE 9. Reproductive morphology of H. okinawa
(CASIZ 144093). Abbreviations: am = ampulla, be = bursa
copulatrix, dd = deferent duct, fgm = female gland mass, ga
= genital atrium, p = penis, rs = receptaculum seminis, v =
vagina. Scale = 0.5 mm.
between the recent specimens examined (Figs.
11A—D) and in both the original description
(Eliot, 1904) and in Rudman’s (1978) descrip-
tion.
REPRODUCTIVE SYSTEM.— The reproduc-
tive morphology of the specimens examined for
this study (Fig. 9) match the description provid-
ed by Rudman (1978).
REMARKS.— Halgerda willeyi, at first
glance, most closely resembles H. iota in exter-
nal morphology, at least based on the original
description of H. iota (Yonow 1993). The phy-
logenetic analysis of Halgerda (Fahey and
Gosliner 2001) supports a close, morphologi-
cally indistinguishable relationship.
Regrettfully, the reproductive anatomy of H.
FiGure 10. Color variation of Halgerda willeyi. A. Jervis
Bay, NSW, Australia; photographed by L. Wiseman. B.
Hachijo Island, Japan; photographed by N. Masatoshi. C.
Okinawa; photographed by R. Bolland. D. Lord Howe
Island; photographed by Ian Hutton.
404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
SOkY Woe Wmm File Name = Ma_inner-20.07
EE, es
m. B. Outer lat-
eral teeth. Scale = 10 um. C. Inner lateral teeth. Scale = 20 um. D. Middle lateral teeth. Scale = 30 um.
iota is unknown, and until additional specimens of H. iota are collected and the internal anatomy
examined, no further comparison of H. iota to other species is possible.
Halgerda willeyi also is externally similar to other Halgerda species. For instance, Carlson and
Hoff compared H. willeyi to H. okinawa and to H. johnsonorum (see Carlson and Hoff 2000). There
are also some external similarities between H. willeyi and H. elegans (see Fig. 7C [from Okinawa
SlugSite and SeaSlug Forum; photo by B. Picton, SeaSlug Forum, March 8, 2000]); both species
have dark lines perpendicular to the mantle edge and yellow-orange lines along the ridge crests.
However, the dark marginal lines of H. elegans do not extend to as large a degree from the mantle
edge throughout the dorsum and into the ridge concavities as they do with H. willeyi. The dark lines
on specimens of H. willeyi are much more numerous, with complex patterns that merge and inter-
weave with the yellow or orange lines all over the dorsum. The yellow ridge crests are the most
notable feature of H. elegans whereas it is the dark lines with the orange or yellow ridges that are
most notable on specimens of H. willeyi.
The gill in these two species is quite different as well. In H. elegans, it is sparse, irregularly
pinnate and has dark coloration encircling the top half of each of the leaves; in H. willeyi, is quite
bushy, feathered and has a dark line extending the length of each of the leaves.
The rhinophores of the two species also differ in coloration. Those of H. willeyi have dark
stripes, those of H. elegans have black subapical coloration.
The reproductive morphology is vastly different between Halgerda willeyi and H. elegans. The
FAHEY AND GOSLINER: HALGERDA GRAPHICA 405
most obvious differences are that H. willeyi has
a large, bulbous penial sheath and a muscular-
ized vagina, whereas H. elegans has a tubular
penis and a glandular vagina.
Other conspicuous internal differences are
noted with the radular morphology of the two
species. Halgerda elegans has fimbriate outer
lateral teeth, with some specimens having mul-
tifid teeth with or without pronounced flanges
(Bergh 1905; Gosliner and Fahey 1998). The
teeth of H. willeyi are simple, hamate and the
outer two or three are flattened plates (Rudman
1978; Carlson and Hoff 2000).
Although color patterns can vary enor-
mously within Halgerda species, as apparent
from recently published photographs cited
herein, field identifications can be assisted by FicuRE 12. Reproductive morphology of H. willeyi
consideration of not just one or two characters, (CASIZ 144095). Abbreviations: am = ampulla, be = bursa
aes copulatrix, dd = deferent duct, fgm = female gland mass, ga
taken by themselves, but by the combination of = genital atrium, p = penis, rs = receptaculum seminis, v =
characters observed. Examination of the inter- vagina. Scale = 0.8 mm.
nal morphology will probably be necessary to
confirm the more difficult-to-identify specimens.
ACKNOWLEDGEMENTS
The authors are grateful to the following individuals for providing the specimens used for this
study: The South Australian Museum; Bob Bolland, Okinawa; Valda Fraser, South Africa and Rie
Nakano, Japan. The Queensland Museum, in particular John Hooper, facilitated this work by mak-
ing the Museum’s facilities and resources available. The authors are further indebted to those pho-
tographers who generously allowed us to use their photos in this publication (their names are asso-
ciated with the individual images).
Financial support for this project was provided by the National Science Foundation PEET
grant DEB 9978155, “Phylogenetic Systematics of Dorid Nudibranchs” to T.M. Gosliner.
LITERATURE CITED
BASEDOW, H., AND C. HEDLEY. 1905. South Australian Nudibranchs, and an enumeration of the known
Australian species. Transactions of the Royal Society of South Australia 20:34—60.
BeERGH, R. 1880. Beitrage zur Kenntniss der japanischen nudibranchien. I. Verhandlungen der koniglich-
kaiserlichen Zoologisch-botanischen Gesellschaft in Wein 30:155—200.
BeERGH, R. 1905. Die Opisthobranchiata der Siboga-Expedition. Monographie. 1—248, pls. 1-20 pp.
BOLLAND, R. 2003. Okinawa SlugSite, vol. 2003. http://rfbolland.com/okislugs/index.html.
CARLSON, C., AND P.J. Horr. 1973. Three new Halgerda species. (Doridoidea: Nudibranchia: Opistho-
branchia) from Guam. The Veliger 36(1):16-26.
CARLSON, C., AND P.J. Horr. 2000. Three new Pacific species of Halgerda (Opisthobranchia: Nudibranchia:
Doridoidea). The Veliger 43(2):154-163.
COLEMAN, N. 1975. What shell is that? Paul Hamlyn Pty. Limited, Sydney. Australia. 308 pp.
COLEMAN, N. 2001. 1001 Nudibranchs. Neville Coleman's Underwater Geographic Pty. Limited, Springwood.
144 pp.
406 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 23
DeBELIUS, H. 1998. Nudibranchs and Sea Snails: Indo-Pacific Field Guide, 2 edition. IKAN-Unterwasser-
archiv, Frankfurt, Germany. 321 pp.
E.tot, C. 1904. On some nudibranchs from East Africa and Zanzibar. Part III. Proceedings of the Zoological
Society of London 1903(2):354—-385.
FAHEY, S.J., AND T.M. GOSLINER. 1999a. Description of three new species of Halgerda from the Western
Indian Ocean with a redescription of Halgerda formosa, Bergh, 1880. Proceedings of the California
Academy of Sciences 51(8):365-383.
FAHEY, S.J., AND T.M. GOSLINER. 1999b. Preliminary phylogeny of Halgerda (Nudibranchia: Halgerdidae)
from the tropical Indo-Pacific with descriptions of three new species. Proceedings of the California
Academy of Sciences 51(11):425—448.
FAHEY, S.J., AND T.M. GOSLINER. 2000. New records of Halgerda Bergh, 1880 (Opisthobranchia: Nudi-
branchia) from the deep Western Pacific Ocean, with descriptions of four new species. Zoosystema
22(3):471-498.
Fauey, S.J., AND T.M. GOsLINER. 2001a. On the genus Halgerda (Nudibranchia: Halgerdidae) from Western
Australia with descriptions of four new species. Bollettino Malacologico 37(5—8):55—76.
FAHEY, S.J., AND T.M. GOSLINER. 2001b. The phylogeny of Halgerda (Opisthobranchia, Nudibranchia) with
the description of a new species from Okinawa. Zoologica Scripta 30(3):199-213.
GOSLINER, T.M., D.W. BEHRENS AND G.C. WILLIAMS. 1996. Coral Reef Animals of the Indo-Pacific: Animal
Life from Africa to Hawaii, Exclusive of the Vertebrates. Sea Challengers, Monterey, California. 314 pp.
GOSLINER, T.M., AND S.J. FAHEY. 1998. Description of a new species of Halgerda from the Indo-Pacific with
a redescription of Halgerda elegans Bergh, 1905. Proceedings of the California Academy of Sciences
50(15):347-359.
Kay, E.A. 1979. Hawaiian Marine Shells. Special Publication edition. Bernice P. Bishop Museum, Honolulu.
653 pp.
Kay, E.A., AND D.K. YOUNG. 1969. The Doridacea (Opisthobranchia: Mollusca) of the Hawaiian Islands.
Pacific Science 23:172-231.
MARSHALL, J.G. AND R.C. WILLAN. 1999. Nudibranchs of Heron Island, Great Barrier Reef. Backhuys,
Leiden. 257 pp.
Ono, A. 1999. Opisthobranchs of Kerama Islands. TBS-Britannica Co., Ltd., Tokyo. 183 pp.
RUDMAN, W.B. 1978. The dorid opisthobranch genera Halgerda Bergh and Sclerodoris Eliot from the Indo-
West Pacific. Zoological Journal of the Linnean Society, London 68:59-87.
RUDMAN, W.B. 2003. SeaSlug Forum, vol. 2003. http://www.seaslugforum.net.
WELLS, F.E., AND C.W. Bryce. 1993. Sea Slugs of Western Australia. Western Australian Museum, Perth. 184
Pp:
WILLAN, R.C., AND G.D. Bropig. 1989. The nudibranch Halgerda aurantiomaculata (Allan, 1932) (Dorid-
oidea: Dorididae) in Fijian waters. The Veliger 32(1):69-80.
Yonow, N. 1993. Opisthobranchs from the Maldive Islands, including descriptions of seven new species
(Mollusca: Gastropoda). Revue Francaise d'Aquariologie 20(4):97-130.
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 24, pp. 407-462, 80 figs. November 14, 2003
The Dangerously Venomous Snakes of Myanmar
Illustrated Checklist with Keys
Alan E. Leviton!, Guinevere O.U. Wogan!, Michelle S. Koo!,
George R. Zug”, Rhonda S. Lucas!, and Jens V. Vindum!
! California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118;
2 National Museum of Natural History, Smithsonian Institution, Washington, DC 20560
No fewer than 39 species of dangerously venomous snakes are currently known to inhabit
Myanmar and the adjacent coastal waters. Of these, 15 are sea snakes and except for two,
Laticauda colubrina and Laticauda laticauda, none voluntarily come onto land (occasionally, obli-
gate marine species may be carried onto shore during severe weather by wave action or enter river
deltas in brackish water). Of the remaining 24 species, all are terrestrial in the sense that none vol-
untarily enters coastal waters. And, of the terrestrial forms, several, such as Trimeresurus albo-
labris, favor arboreal habitats and are usually found resting on tree limbs. All of the terrestrial
species can swim, and some are occasionally found swimming in the rivers and streams as well as
in flooded rice paddies.
Two families of dangerously venomous snakes are represented in the Myanmar herpetofauna:
Elapidae (cobras, kraits, and coral snakes [subfamily Elapinae], and sea snakes and Australian
elapids [subfamily Hydrophiinae]), and Viperidae (true vipers [subfamily Viperinae], pitvipers
[subfamily Crotalinae], and Azemiops [subfamily Azemiopinae]). Known mildly venomous
snakes found there, mostly referred to the very large snake family Colubridae, include the rear-
fanged snakes of the Asian vine or whip snake genus Ahaetulla, the cat-eyed snakes (genus Boiga),
the genus Psammodynastes, and the aquatic and semi-aquatic snakes of the genera Enhydris,
Cerberus, Cantoria, Fordonia, and Bitia. How dangerous any of these are to humans is still an
open matter for research. In a like manner, several of the supposedly nonvenomous colubrids have
been shown to have toxic salivas and some should be considered mildly if not dangerously ven-
omous. Among these are members of the genera Xenochrophis, Amphiesma, and Rhabdophis. What
we do know is that we often underestimate the severity of many snakebites of both juvenile dan-
gerously venomous and supposedly non-dangerously venomous snakes. Yet, bites of just such ani-
mals have been implicated in the deaths of several well-known and knowledgeable professional
herpetologists, notably Karl Patterson Schmidt who, in 1957, at the age of 67, was bitten by a juve-
nile boomslang (Dispholidus typus), an African rear-fanged snake, and died one day later, Fred
Shannon, M.D., who, in 1965, was bitten by Crotalus scutellatus, and died shortly thereafter,
Robert Mertens, who was bitten by the African rear-fanged snake, Thelotornis capensis, in 1975,
and most recently, Joseph Slowinski, who at the age of 38 was bitten by a 30-cm long juvenile krait,
Bungarus sp.*, and died within 48 hours. (See Appendix A for comments relating to procedures for
handling venomous snakebites in Myanmar.)
Needless to say, care should always be exercised when handling any snake, even those that are
supposedly nonvenomous. And, it must be remembered, it is often difficult to distinguish ven-
omous and nonvenomous species without careful inspection. In Myanmar, for instance, several
3 Ulrich Kuch (Universitat Frankfurt) believes that the specimen, although closely resembling Bungarus wanghaotin-
gi, represents a distinct species, which he is in the process of describing (see also remarks under Bungarus wanghaotingi).
407
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
408
Volume 54, No. 24
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© 2003 California Academy of Sciences
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ical map of Myanmar
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LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 409
snakes are black with white bandings; among these are Lycodon zawi, Lycodon fasciatus, Dinodon
septentrionalis, Bungarus wanghaotingi, and Bungarus flaviceps. The first three are nonvenomous,
the latter two dangerously venomous. Although the five can be told apart fairly readily on the basis
of color patterns, once one knows the differences, yet, the only sure way to know whether the snake
is a venomous or nonvenomous species is to look at the side of the head just in front of the eye to
determine if a loreal scale is present or not. In the illustrated keys that follow, these and other defin-
ing characters are clearly noted.
The varied topography and associated ecozones of Myanmar (see Maps p. 408 and 458) sup-
port a highly diverse herpetofauna, and we are just now beginning to learn how rich in species it
really is. The country has not been carefully studied for many years, and it is a credit to the goy-
ernment of Myanmar and especially to the past and present directors of the Nature and Wildlife
Conservation Division, Forest Department, U Uga and U Khin Maung Zaw, that intensive surveys
are being conducted throughout the country. Already, significant results have been achieved.
Shortly before his death, Joseph Slowinski and his colleague Wolfgang Wiister determined that a
hooded cobra found in the Mandalay region represented a distinct species, which they named Naja
mandalayensis. The importance of this can be quickly appreciated when it is understood that to
treat snakebites one most often uses species-specific antivenoms, species-specific in that the
antivenom used to treat one kind of snakebite usually will not work for the bites of other species.
In Myanmar, for instance, only Naja kaouthia antivenom is available for cobra bites. Whether this
species-specific antivenom works for other cobra snakebites is unknown. Inasmuch as N. man-
dalayensis was only recently recognized as distinct from N. kaouthia, there 1s no way of telling how
many people, bitten by N. mandalayensis but treated with N. kaouthia antivenom, died but could
have been saved had more been known about the species diversity and had the proper antivenom
been available. Thus, it is hoped that this contribution will enable those engaged in the study of the
fauna in the field and the laboratory to recognize more readily the most dangerous as well as inter-
esting components of that fauna.
In the following checklist, we have made no attempt to provide inclusive synonymies. They
are available in the works we do cite, namely, Smith (1926 and 1943), Golay et al., especially the
sections by McCarthy and by Toribe (1993), Bauer (1998), McDiarmid, Campbell and Touré
(1999) and David and Ineich (1999). Where necessary, we do provide additional references. In the
statements on distribution, again we have kept them brief, confining ourselves to country for non-
Myanmar localities and, where known, to State or Division within Myanmar.
With respect to species names, we have accepted names for several of the dangerously ven-
omously snakes in Myanmar that have not been widely used in the earlier literature. One notable
instance is the many-banded krait, Bungarus wanghaotingi, formerly Bungarus multicinctus.
Recent work indicates that the former is both morphologically and geographically distinct from the
latter and is the species occurring in Myanmar (but see footnote 3) and the neighboring region of
Yunnan Province, China. Bungarus multicinctus, in its restricted sense, is known from eastern
China, Taiwan, and to the south and west to Laos. In a like manner, we recognize Trimeresurus yun-
nanensis as distinct from T: stejnegeri and, tentatively, have removed the latter from the faunal list
for Myanmar. David et al. (2001:219) posit that 7: stejnegeri (sensu stricto) may yet be found in
eastern and possibly northern Myanmar (T. stejnegeri is known from China [including Yunnan
Province], Taiwan, Vietnam, and Laos [see David 2001:218]). If so, we suspect that almost certain-
ly it will be found in northeastern Shan State inasmuch as it has been taken in the vicinity of
Menglian, in southwestern Yunnan, not far from the border with Myanmar.
A word about treating snakebites. First, all bites should be taken seriously. It is true that a large
percentage of the bites, even by dangerously venomous snakes, are what are known as “dry bites,”
410 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 24
that is bites in which no envenomation takes place. But, it is not always possible to know this in
advance. So, best take no chances and seek appropriate medical treatment immediately. Following
the section on the identification of dangerously venomous snakes, we have appended a brief state-
ment on how best to proceed in the event of a snakebite. Although more complete instructions are
available elsewhere and reference to several published articles is given in the bibliography, we have
included as Appendix A a brief statement on the treatment of snakebite in Myanmar prepared in
2000 by Dr. Slowinski for use by members of the Myanmar Herpetological Survey field teams.
This publication was prepared initially for distribution within the Nature and Wildlife
Conservation Division of the Forest Department, Ministry of Forestry, Myanmar, and specifically
to members of the Myanmar Herpetological Survey field team in Myanmar. First written in April
2002, it has been revised to include new data gathered both by the survey’s field team and by the
authors’ reexamination of both specimens and literature. However, there are a number of people in
laboratories in Europe and the United States who are actively engaged in research dealing with the
viperid genera Trimeresurus, Ovophis, and Protobothrops, the snakes of the genus Bungarus, and
the sea snakes, so that the scheme of classification of the venomous snakes as presented here will
likely change in the near future.
ACKNOWLEDGMENTS
The authors would like to take this opportunity to express their profound appreciation to U
Shwe Kyaw, Director General, Forest Department, Ministry of Forestry, and U Khin Maung Zaw,
Director of the Division of Nature and Wildlife Conservation, Myanmar, for their ongoing support
and encouragement of the Myanmar Herpetological Survey. The survey, initiated by Joseph Bruno
Slowinski, Assistant Curator of Herpetology, California Academy of Sciences, in 1998, was initial-
ly conceived by U Uga, then Director of the Nature and Wildlife Conservation Division (NWCD).
It was with U Uga’s encouragement and endorsement that Dr. Slowinski and his colleague, Dr.
George R. Zug, Curator of Amphibians and Reptiles at the Smithsonian Institution’s National
Museum of Natural History, applied for and were granted substantial financial support by the
National Science Foundation. The survey was begun in earnest in 1999 at about the same time that
U Khin Maung Zaw succeeded U Uga as Director of the NWCD. U Khin Maung Zaw, without hes-
itation, confirmed his division’s confidence in the project. With this support, and with the assign-
ment of several members of the NWCD to the project on a continuing basis, Dr. Slowinski, and
other members of the Academy’s staff, including Dong Lin, staff photographer, Jens Vindum,
Senior Collections Manager in the Department of Herpetology, and two graduate students in
Herpetology, Ms. Rhonda Lucas and Ms. Guin Wogan, traveled to Myanmar to conduct field work
and, as part of their commitment to the NWCD, to assist in the training of members of its staff in
both field and museum techniques.
As noted earlier, in September of 2001, while doing field work in northern Kachin State, Dr.
Slowinski was bitten by a krait, Bungarus sp., that had just been collected. Because of the field
party's isolation, it was not possible to obtain medical help in time and despite valiant efforts to
keep him alive, on 12 September 2001, Dr. Slowinski died.
Following the loss of Dr. Slowinski, it was decided by members of the staff of the Academy’s
Department of Herpetology, again with the encouragement of U Khin Maung Zaw, to continue the
project. Thus, we take this opportunity to acknowledge with thanks the contributions that have been
and are currently being made to this work by a group of dedicated people both in Myanmar and in
the United States: in Myanmar — U Shwe Kyaw, U Khin Maung Zaw, Director of the Division of
Nature and Wildlife Conservation; NWCD staff: U Htun Win, Daw Thin Thin, U San Lwin Oo, Sai
Wunna Kyi, U Kyi Soe Lwin, U Awan Khwi Shien, and U Hla Tun; in the United States: the late
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 411
Joseph Bruno Slowinski, Dong Lin, and Douglas Long. Special thanks must also be accorded U
Hla Tun, Dong Lin, Francis Lim, Ashok Captain, Indraneil Das, Nikolai Orlov, and John Tashjian
who are responsible for many of the excellent photographs of venomous snakes that are reproduced
here. The computer-generated distribution maps were prepared by Michelle S. Koo, the
Department of Herpetology’s resident Biogeographical Information System Coordinator.
Dr. Indraneil Das, to whom we are indebted for his critical review of the manuscript, also pro-
vided us with extensive new information that he has garnered from several sources, including
Bauer (1998), McCarthy (1993), Toriba (1993), and his own researches, that either correct or
amplify what is known about type localities and the location of holotypes or syntypes of many of
the species we deal with here: Bungarus bungaroides, Bungarus fasciatus, Laticauda colubrina,
Laticauda laticaudata, Naja kaouthia, Ophiophagus hannah, Sinomicrurus macclellandi,
Enhydrina schistosa, Hydrophis cantoris, Hydrophis fasciatus, Hydrophis gracilis, Hydrophis
obscurus, Hydrophis ornatus, Hydrophis spiralis, Hydrophis stricticollis, Kerilia jerdoni,
Praescutata viperina, Daboia russelii, Ovophis monticola, Protobothrops jerdonii, Protobothrops
mucrosquamatus, Trimeresurus erythrurus, Trimeresurus popeiorum, and Trimeresurus stejnegeri.
Dr. Das offered that we could include his data here, but we have chosen not to do so because short-
ly he will be issuing an updated checklist of Indian reptiles and will include these data in that work.
We do want to express our deepest appreciation for his generosity and now look forward, eagerly,
to his publication.
At this time we also want to acknowledge ERSI’s Conservation Technology Support Program
(CTSP) for generously providing both the software and training that have enabled us to prepare the
distribution maps that accompany this report.
Lastly, we want to express our appreciation to Dr. Michele L. Aldrich who, with her usual edi-
torial acumen, read the manuscript and caught more errors than we wish to acknowledge.
The Myanmar work has been supported by a grant from the National Science Foundation,
DEB 9971861. Additional funding has been provided by the California Academy of Sciences’
Research Division Inhouse Research and Geraldine K. Lindsay Funds and the Department of
Herpetology’s Dufflebag Fund.
This contribution is dedicated to the
memory of
Dr. Joseph Bruno Slowinski
who lost his life in the pursuit of an
understanding of the Myanmar
herpetofauna, with special reference to
its dangerously venomous snakes.
412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 24
oe revs pee
EEO
r labia eH : ee
Soden ocen oC
stahebeataeeate
ia
aren ca
£\
ody scales (numbers indicate best hod for coun scale
Scales arranged in (A) oblique rows; (B) parallel rows
(Modified from Smith [1943])
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 413
CHECKLIST AND KEYS TO THE DANGEROUSLY VENOMOUS SNAKES OF MYANMAR
Key to the Genera and Species of Venomous Snakes of Myanmar!
la. Pupil of eye round; loreal scale absent (Fig. 1); poison fangs short, permanently erect; maxil-
lary bone elongate, usually with several small teeth behind the front fangs (Family Elapidae,
including cobras, coral snakes, and sea snakes)
1b. Pupil of eye vertically elliptical; loreal scale present (Fig. 2) or, if absent, then upper surface
of head either covered by small scales (Fig. 3) or, if covered by large, symmetrical shields,
then a deep sensory pit present immediately behind the nostril (Fig. 4) (also present in many
in which the head is covered by small scales); poison fangs variable in length, fixed to a short
moveable maxillary bone that allows for rotation of the fangs backward when the mouth is
closed no) post-fane teeth: behind tangs (Family Viperidae)- 4. 224-4. eee jay)
Oe 3. ae ee
Left to right: (1) L otingi); (2) loreal present, separating nasal
and preocular (Azemiops feae); (3) head covered with small scales, sensory pit absent (Daboia russelii);
(4) head covered with small scales, sensory pit present (7rimeresurus erythrurus)
2a. Tail flattened laterally, oar-like (Fig. 5)
PAeeLACECOUNGEM apenine (EIR 6) ey eaters ein mene meremae tore boat caus etic epee rtcmlnns, Sue mente Urey th 8)
(5) Oar-like tail (Laticauda laticaudata), which is characteristic of all venomous sea snakes versus
(6) a tapering or at least non-flattened tail (Trimeresurus purpureomaculatus), characteristic of all terrestrial snakes
3a. Vertebral series of scales not enlarged (Fig. 7); scales on sides of body obliquely arranged or
| 0] Ei as cae Ae race Ae Pr ea Mere tr PMG nt a MMP ee ery Et Lee Ueee hae oC 4
3b. Vertebral series of scales enlarged (Fig. 8), distinctly larger than scales on sides of body; scales
on sidesiot body motobliquelysamancediyer,.. 5 2 ace eer ie aera ©: 9
! In preparing this key and the checklist that follows, we have been guided by the latest revisionary studies, but we
have also taken a conservative position because of the nature of the animals with which we are working. Thus, although we
accept McDowell’s argument that among the sea snakes Enhydrina schistosa should be referred to the genus Disteira and
Thalassophis viperina to Lapemis, we have not done so here. Most of the medical and general literature dealing with sea
snakes still refer to Enhydrina schistosa and Thalassophina viperina. Also, we have, rather arbitrarily, chosen to accord all
recognizably distinct allopatric subspecies full species status rather than get embroiled in endless arguments of why raise
one subspecies to full species status but retain another as a subspecies, though both can be readily, though differentially,
diagnosed taxonomically. Arguments as to what constititues a biological “species” versus “subspecies” are pointless and are
usually based on an arbitrary assessment by a given worker on just how “important” a given character or character state is
in inferring the “closeness” of biological affinities, i.e. the genealogical relationships among the parties.
414 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 24
4a. Scales on sides of body obliquely arranged (Fig. 9), pointing backward and downward, most
distinct on sides of neck; scales in 15 to 23 longitudinal rows around midbody .......... 5)
4b. Scales on sides of body not obliquely arranged (Fig. 10), in straight longitudinal rows; scales
il Satoml sslongitudinalirowsraround midbodyemeere ass cse. saeaaees ebb oo 82g eccc ee 7
- — - B Bs "4 2s ee & - ee = Led : ph i
Left to right: (1) Vertebral row of scales not enlarged, (2) vertebral scales distinctly enlarged (Bungarus wanghaotingi);
(3) body scales, except for outer two or three horizontal rows, obliquely arranged (Naja mandalayensis);
(4) body scales in parallel rows, not obliquely arranged
5a. Scales in 15 longitudinal rows at midbody; a pair of large occipital shields present (Fig. 11).
5S ES ETE ENCE OT RRR Tet as SL oe eS Ophiophagus hannah
5b. Scales in 19-25 longitudinal rows at midbody; no enlarged occipital shields (Fig. 12)..... 6
Occipital d
scales small
Y | atetetige,
large ‘ ,
11 Scales larg : > . ; 12
Head scutellation in cobras: (11) Paired occipital shields present (Ophiophagus hannah);
(12) No enlarged occipital shields (Naja sp.)
6a. Usually distinct dark markings on the hood; throat cream colored, often with a pair of well-
defined lateral spots; when present, only a single dark band on the throat, otherwise venter
either pale or gradually increasingly cloudy with dark pigment towards the rear, posteriorly
venter Oftenrtotally darks. 2e..)s8.5% .6./> «io See ee ee Naja kaouthia
6b. No or only faintly distinct markings on the hood; throat extensively darkly mottled anterior to
first dark throat band, mottling obscuring the throat spots characteristic of most species of
Naja; at least two distinct dark bands on venter, one on the throat followed by a second on
anterior third of venter; venter otherwise pale with some dark mottling Naja mandalayensis
Tas nal scute. divided: (Fig. 13) sc. cc Sic de since = Spe oe a ie Soe an 8
7b. Anal scute single (Fig. 14); preocular scale present; snout and venter free of dark pigment (no
GME daerecorasion Myanmar). <4... «| ..6/ see ya cede nine eee eee ene Calliophis bivirgatus
undivided anal scute
B divided anal scute 14
(13) Anal plate divided (Sinomicrurus macclellandi);
(14) Anal plate undivided (single) (Calliophis bivirgata)
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 415
8a. Pattern on dorsum characterized by small black spots (Fig. 15), sometimes forming longitudi-
nal stripes, with two black bands or rings on tail, one at base and one at the tip; one to three
smallimaxillanyiteethi behind fangsia. 25,9 ase aeies: ieee canine = Calliophis maculiceps
8b. Pattern on dorsum characterized by small but distinct black vertical bars on sides (Fig. 16),
sometimes forming crossbars, with two black rings on tail; no maxillary teeth behind fangs
5 ER Rye SPC. ROLES A eM MENTE REET. CEN aT ce Ge Sinomicrurus macclellandi
Dorsal color patterns: (15) Spotted pattern of Calliophis maculiceps;
(16) Barred pattern of Sinomicrurus macclellandi
9a. Scales in 13 longitudinal rows at midbody; terminal caudal scales paired
MPP ry niet arora acs Wiaiera Khe shall lmete kts @ etn tieral o Me aarp Bungarus flaviceps
9b. Scales in 15 longitudinal rows at midbody; terminal caudal scales single or paired ...... 10
10a. Subcaudal scales usually paired, occasionally some unpaired anteriorly (Fig. 17), but terminal
SCaleSpaliwayGapalne Gs tei ei2y sarin, autsier eyaira gies bag Gaeeeese Mellons) Hereralaice lee DORE Bungarus bungaroides
fobs Subcaudaliscales entire (Eig 41/8) smot pained saan ewe ome ele oye ene il
fees Anterior
18
Subcaudal scutes: (17) Mixed paired and unpaired (Bungarus bungaroides);
(18) All single (unpaired) (Bungarus fasciatus)
lla. Tail tapers to a point (Fig. 6); neural processes of vertebrae not enlarged vertically and do not
produce a distinct ridge down the back; dorsal crossbars do not encircle body; belly white 12
11b. Tail ends bluntly, not pointed; neural processes of vertebrae enlarged vertically and produce a
distinct ridge down the back; body marked with alternating black and yellow bands that com-
pletelygzencincleDOUY ci. a= ope ohomumor tenets Stet ek EUS Re rs ie Bungarus fasciatus
12a. 27-48 narrow white crossbars on back, crossbars narrowest middorsally, expanding on sides
PR ns ee OR Tee elses Ale ae ake eae Ako acsebeeetinien eeeee egal ape remany ee Bungarus wanghaotingi
12b. 11—14 broad white, black-spotted crossbars on back, crossbars as broad as or broader than the
MatkeimIntCISPACESi reat ney eer hee «, « See aE an eee Bungarus magnimaculatus
13a. Ventral scales large, one-third to one-half the width of the body; maxillary bone extends for-
wards beyond palatines; nostrils lateral, nasal scales separated by internasals........... 14
13b. Ventral scales small, less than one-fourth width of body, often smaller than or at least not larg-
er than adjacent body scales; maxillary bone usually does not extend forwards beyond palatine
(exceptions, Kerilia jerdoni, Hydrophis gracilis and H. cantoris); nostrils variable, internasal
scales ausent nasal scalesm contact wath’ one another ya. ser eic a ae meee alors 5)
416 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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14a. Scales in 19 longitudinal rows at midbody; no azygous (median) prefrontal scale (Fig. 19)
ree: OSA Pes te Sls eld Jy oS nd el teen Re eis Laticauda laticaudata
14b. Scales in 21—25 longitudinal rows at midbody; an azygous (median) prefrontal scale usually
Presents (os 2()) ett sad moet ns cuces tevtd cteees AS eeeG See ee ee Laticauda colubrina
20
~ No azygous prefrontal; i, Azygous prefrontal
prefrontals in contact separates prefrontals
Azygous prefrontal shield: (19) Absent in Laticauda laticaudata;
(20) Present in Laticauda colubrina
15a. Ventral scales small but distinct, undivided by a median groove or, if divided posteriorly, the
two halves either apposed or alternating with one another, then head very small and body long
andivery Slender anteriorly... 7226 Sia). 1c «os eee eae ones se ne 16
15b. Ventral scales, if distinct, then either divided by a median groove or smaller than adjacent
Lo Yer hii er: | (tee ee ne aes cele ohh Sree hts olats 1 CMe Ce eran oS 0100 00 00 28
16a. Scale rows in 23 or fewer rows around midbody; maxillary bone extends forwards beyond
palatine; little or no space (diastema) separates fangs from smaller maxillary teeth
RI ai Pee Me Penn ane nt reee ee TI EMME ee RR Eris cies SS ERY ce Gino oc Kerilia jerdoni
16b. Scale rows in 25 or more rows around midbody (but often fewer around neck); maxillary bone
does not extend forwards beyond palatine (except in H. gracilis and H. cantoris); a distinct
space (diastema) separates fangs from smaller maxillary teeth ...................... 17
ivia-dMentalyscalé:normali (Pigs 21): tis cpheeiotealt Aes ane ae AR eee 18
17b. Mental scale elongate, partially hidden in groove in the symphysis (Fig. 22); ventrals uniform
imisize+5—9) smallimaxillary teeth behind fangs 2%... f.0 s5 -- seca eee Enhydrina schistosa
Mental shield normal, not Mental shield elongate,
elongate, not in groove in : inserted in groove in
symphysis = symphysis
Mental shield: (21) Mental normal, neither elongate nor partially hidden in groove in the symphysis (Hydrophis);
(22) Mental elongate, hidden in groove of symphysis when mouth is closed (Enhydrina schistosa)
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 417
18a. Ventral scales broad anteriorly, reduced posteriorly and not distinct from adjacent scales (Fig.
2) sounanillanytecthibelind tangs... 5 9c os oe eee See ee Thalassophina viperina
18b. Ventral scales slightly distinct from adjacent scales and of uniform shape throughout irrespec-
tive of their size, (Fig. 24); 1-18 maxillary teeth behind fangs ...................... ng
Ventral scutes: (23) Undivided and indistinguishable from adjacent scales (Lapemis hardwickii); (24) Usually distinguishable
from adjacent scales (Hydrophis cyanocinctus); (25) Divided by a longitudinal groove (Hydrophis gracilis)
(From Smith, 1926, pl. 1, figs. 3, 4, and 1.)
19a. Head very small, body elongate, anterior fifth slender, about same diameter as head (Fig. 27);
ventrals small, posterior series usually divided by a longitudinal groove (Fig. 25), the two
halves either apposed or alternating with one another; maxillary bone extends as far forward
as anterior tip of palatine or beyond; 5—6 small maxillary teeth behind anterior fangs .. . . 20
19b. Head not distinctly reduced, body not particularly slender anteriorly (Fig. 26); ventrals small,
usually distinct throughout and entire, only rarely are a few divided by a longitudinal groove;
maxillary bone does not extend forward beyond palatine; 1-18 small maxillary teeth behind
ACER OL AM OS hc et eile sce oe Ee tee Cee TET Re ey A da CR Ie LAE SARE 4 Steir Ze
Head and body dimensions: (26) head and anterior portion of body not distinctly narrowed (Hydrophis spiralis),
(27) head and anterior fourth of body slender (Hydrophis atriceps)
ie emitalscales entire throushout: head black: semi 2 eo on ee en eee 21
20b. Ventral scales anteriorly entire, posteriorly at least some divided by a longitudinal groove
(Eile) sheadieOlOrevablaDles, seri. s «oscil sea ere 6 here wus Sve eae ane te SE alice Pe ecao nes 22
21a. Scales in 28-33 rows on neck, 49-58 around midbody; ventrals 414-514
TS Lee ae PORN Ne AE sat ta Tree Myc tito. sR ay, Sok a hai Hydrophis (Hydrophis) fasciatus
21b. Scales in 25-30 rows on neck, 39-49 around midbody; ventrals 323-452
eA NE ee a ae, MERA TAS Re Le: HAGE, Hydrophis (Hydrophis) atriceps
418 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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22a. Prefrontal scale usually in contact with second upper labial (Fig. 28); ventrals 220-350; 17-23
ScalesmonvSearounGd Meck eo = tae ae See oe Hydrophis (Hydrophis) gracilis
22b. Prefrontal scale usually in contact with third upper labial; ventrals 404-468; 23-25 scale rows
ATOUNd Ee ckesrane ly lea. seein eee ees PS Wee Hydrophis (Hydrophis) cantoris
Prefrontal
28
(28) Lateral view of head of Hydrophis gracilis showing contact of prefrontal
with second upper labial; third and fourth upper labials border eye
Second upper labial
23a. Scales at midbody rounded or bluntly pointed posteriorly, feebly to distinctly imbricate; 1—8
maxillary teeth behind anteniortangs, 2202 Noe. Sele Ie eee Se ce eee 24
23b. Scales at midbody feebly imbricate or juxtaposed, quadrangular to hexagonal in shape; 8-18
maxillary teeth’ behind anterionfangs 42.49.4226 ae sents oe ie a ee ee 26
24a. No enlarged anterior temporal scute, temporals small, scarcely distinguishable from surround-
ing scales; 1-2 maxillary teeth behind anterior fangs................ Disteira nigrocincta
24b. Normally one large anterior temporal scute (Fig. 29), clearly distinguishable from ordinarily
scales, occasionally extending downward to border of lip.................2.2000 00s 25
25a. 19-23 scale rows on neck; ventrals 300-338............ Hydrophis (Hydrophis) obscurus
25b. 25-31 scale rows on neck; ventrals 295-362 ........... Hydrophis (Leioselasma) spiralis
26a. Normally two or three anterior temporals (Fig. 30); ventrals less than 350; head olive or gray
in adult; 40-60 dark crossbands or rhomboidal spots, wider than light interspaces, becoming
obscure with age; below yellowish or whitish) 72. .-7..,52 +... 0. =: > here 27
26b. Normally one anterior temporal (Fig. 29); ventrals 374-452; head dark gray or olive, or, if
blackish dorsally, with yellow mottling on snout and sides; dorsum with 45—65 dark bands,
becoming obscure with age; below pale; dentary teeth 19—22
Cee RS Ee Meare Se adn Boar shan nd. ne SRO Hydrophis (Aturia) stricticollis
Large anterior temporal === Two anterior temporals
29
(29) Single large anterior temporal (Hydrophis spiralis); (30) Two anterior temporals (Hydrophis caerulescens)
27a. Pale gray or olive above to almost white, with broad dark bars or rhomboidal spots separated
by narrow interspaces; yellowish or whitish below; head olive; 8-13 maxillary teeth behind
TT) COG PINS EMRE et eA a ae ac gus sc made Seen eee Hydrophis (Aturia) ornatus
27b. Bluish-gray above, yellowish or whitish below, with 40-60 broad bands about twice as broad
as interspaces; markings indistinct on older individuals, which are almost uniform gray; head
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 419
dark gray to black, occasionally with light curved marking in young; 13-18 maxillary teeth
hehindiiamesney en Cate ee AGG ORS te Hydrophis (Aturia) caerulescens
28a. Olive to gray above, whitish below, 35-55 narrow dark bands, occasionally uniform dark
gray; midventral rows of body scales larger than adjacent lateral and dorsolateral scales; ven-
tral scutes, if discernable, not divided by a longitudinal groove; 3—6 small maxillary teeth
behind fangs; no discernable series of midventral scutes ............. Lapemis hardwickii
28b. Bicolored, ordinarily black above, yellow below, with some variation in which yellow extends
further up on the sides and the black is restricted to a wide middorsal stripe; sometimes black
bars on belly; tail mottled yellow and black; ventral scutes, when distinct, divided by a longi-
tudinalboroove=5—0)maxallaryitecth arin seein eee ae ee nae ie Pelamis platurus
29a) Loreal pit present (Fig.4): head distinctly triangular in’shape.) {2.92.2 2.25....---- Bill
29b. Loreal pit absent (Figs. 2—3); head distinct from neck but variable in shape ........... 30
30a. Loreal scale present but small (Fig. 2); head scutes large, symmetrical; dorsal scales in 17 lon-
siuidinal row sat midbOdy.<ccp or. -eccdia ud cere RO ean ne Ree Azemiops feae
30b. Loreal scale absent (Fig. 3); both top and sides of head covered by small, imbricate, distinct-
ly keeled scales, except for the small, strongly crescentric supernasal, the large undivided
nasal andLO—12upperlabialssiv.5 jp ftain geo ee oe ea Daboia russelii
31a. First upper labial completely separated from nasal by a distinct suture (Fig. 31)........ 32
31b: First upper labial partially or completely fused to nasal (Fig. 32)...........5......:. 39
Nasal and first upper labial scales in Trimeresurus and Ovophis: (31) Nasal completely separated from first upper labial by
suture (Ovophis monticola); (32) Nasal partially or completely fused to first upper labial (Trimeresurus purpureomaculatus)
32a. Body scales in 17—21 (rarely 23) longitudinal rows at midbody ..................... 35
32b. Body scales in 23—27 (rarely 21) longitudinal rows at midbody..................... 33
So Peventralscmore thames cree ii c.d,-00 eyed Peon cna memet ete eon ee eee ee ere 34
33b. Ventrals fewer than 190 (137-176 for Myanmar and adjacent areas of India, Thailand, and
China, 127—144 further east); subocular scales usually fragmented into smaller scales
BE See EV apo TR eis tai ahs Ratal ts Nas Bhs Jon 3 hia = ERE Me eet Se ONE CRC ae Ovophis monticola
34a. 8-10 scales in a line between supraoculars; ventrals 201-212; subcaudals 66—78; outermost
one or two rows of body scales (those bordering ventrals) smooth, all other rows strongly
Cee) (a oar ae eee en eR PE SEER: «cotton 8, Cen ol coe Protobothrops kaulbacki
34b. 14-16 scales in a line between supraoculars; ventrals 200-218; subcaudals 76—91; outermost
rows of body scales (those bordering ventrals) keeled ..... Protobothrops mucrosquamatus
35a. Body scales in 17 longitudinal rows at midbody; ventrals fewer than 150; bicolored ventrolat-
eralesthipe present im bothimales:and females 2% ...2.2 264+. +00 Trimeresurus medoensis
35b. Body scales in 19-21 longitudinal rows at midbody; ventrals more than than 150; ventro-
Pera stcipetviaiiab lew pwn waar tive, ante clei ete sake ocdtere Gears eile, Sadia wh slensele wai Om oiene 36
420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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36a. Dominant background body color green; dorsum of head green..................... 37
36b. Dominant background body color variable, greenish or olive above with series of transverse,
rhomboidal, or irregular reddish brown to black spots or blotches, to entirely black; dorsum of
head black with symmetrically arranged yellow markings; belly yellow with black spotting
3 0 0:0 6 6 Bee GG Rae ae Akal opener Tot creme eee! Ate Protobothrops jerdonii
37a. Scales in 19 (rarely 21) longitudinal rows at midbody and 19 (—21) on neck; ventrals 155-165
(—170); subcaudals (S8—) 61-68 (in Myanmar from Kachin and ?Chin States); ventrolateral
stripe bicolored (orange or brown below, white above) in males, white or absent in females
NP ee eT Beech sens er aN Rie wh ot aaelTe Sis wa ee wears eM Trimeresurus yunnanensis
37b. Scales in 21 longitudinal rows at midbody (21—23 on neck); ventrolateral stripe bicolored
(orange or brown below, white above) in males, bicolored or white only in females ..... 38
38a. Ventrals: males 161-172, females 157—169; subcaudals: males 71-79, females 58-74; hemi-
penes long, slender, extending to level of 20— 25) subcaudal scale, forked at level of fifth
subcaudal scale, without spines (in Myanmar, known from vicinity of Mergui); ventrolateral
stripe bicolored (orange or brown below, white above) in males, white in females
PR eee ne rete SS eee eames Secs Se egele ee vel Are eee sae yes Trimeresurus popeiorum
38b. Ventrals (for Chinese specimens only [after Zhao et al. 1998 and David et al. 2001]): 154-172;
subcaudals: 43-75; hemipenes short, stout, extending to 10‘ subcaudal plate, forked at level
of fifth subcaudal scute, spinose (species not presently known from Myanmar but possibly in
north and east); ventrolateral stripe bicolored (orange or brown below, white above) in males,
bicolored*or white inviemales:,.... cee... See eee. tae Trimeresurus stejnegeri
39a. Scales in 21 (rarely 19) longitudinal rows at midbody; temporal scales smooth or weakly
keeled; dorsum of head uniform green; tail usually not spotted with brown (in juveniles, tip of
Cail MLO WI) he. cc-sete sete chee ooh «eRe a tea eae eons ge Trimeresurus albolabris
39b. Scales 23 or more longitudinal rows at midbody; temporal scales keeled; tail usually spotted
WWUING TOWN cas eee oes nue. en ane cea a cress ec aero eee One inate ae ee eee 40
40a. Head uniform green; body green above, pale green to yellowish below; ventrals: males,
153-174, females, 151—180; subcaudals: males, 62—79, females, 49-61
Sole 8 Gare ROU CSUR ech Ses roar acte tPA I Pada tea acct. Trimeresurus erythrurus
40b. Head and body brown to purplish-brown above, whitish to brown below; ventrals: males,
160-179, females 168-183; subcaudals: males 74-76, females 56-63
=o BSCR cat ea rn a Pe aE GD hy RUA roe Trimeresurus purpureomaculatus
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 42]
CHECKLIST OF THE DANGEROUSLY VENOMOUS SNAKES OF MYANMAR
FAMILY ELAPIDAE
Subfamily ELAPINAE
Genus Bungarus Daudin, 1803
Bungarus Daudin, 1803. (Type species: Bungarus annularis Daudin, 1803 [=Pseudoboa fasciata Schneider,
1801)).
Bungarus bungaroides (Cantor, 1839)
Elaps bungaroides Cantor, 1839:33. (Type locality: Cherra Punghi, Khasi Hills, Meghalaya State, India;
Holotype: BMNH 1946.1.17.91).
Bungarus bungaroides, Smith, 1943:410.— Toriba, 1993:118.— David and Ineich, 1999:66.
DIAGNOSTIC CHARACTERS.— Dorsal
scales in 15 longitudinal rows at midbody;
subcaudal scutes ordinarily divided anteriorly,
but occasionally some scutes may be single,
but always divided near the tip; ventrals
220-237; subcaudals 44-51; dorsum black
with a series of very narrow white to pale yel-
lowish lines or crossbars; on the belly, the light Bungarus bungaroides (from Boulenger 1893, pl. 18, fig. 5).
crossbars widen to form distinct transverse
bars. Total length! 1400 mm; tail length 160 mm (largest male).
DISTRIBUTION.— MYANMAR (Map p. 458): Kachin State. ELSEWHERE: India (Sikkim; Assam
[Khasi Hills]); Cachar.
HasitatT.— Historically recorded in northern Myanmar (Smith 1940), this species has been
documented at elevations of 2040 m (Boulenger 1896:371). In Myanmar, to date, this species has
been found only in the subtropical forests of extreme northern Myanmar.
Bungarus fasciatus (Schneider, 1801)
Pseudoboa fasciatus Schneider, 1801:283 (Type locality: Mansoor, Cottah, Bengal, India; Holotype: based on
Russell, 1796:3 and pl. iii).
Bungarus fasciatus, Smith, 1943:411.— Toriba, 1993:119.— David and Ineich, 1999:68.
DIAGNOSTIC CHARACTERS.— Dorsal
scales in 15 longitudinal rows at midbody; sub-
caudal scutes undivided throughout; middorsal
row of scales (vertebrals) strongly enlarged, as
broad as or broader than long; tail end blunt;
distinct vertebral ridge down the back formed
by the neural processes of the vertebrae; ven-
trals 200-234; subcaudals 23-39; pattern of
black and yellow bands, all of which encircle
body. Total length recorded to 2125 mm, said
to be rare over 1800 mm. (Smith, 1943:412.)
DISTRIBUTION.— MYANMAR: widely dis-
tributed (Ayeyarwady Division, Kachin State,
Body and tail lengths are mostly taken from Smith (1943).
422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Magway Division, Mandalay Division, Rakhine State, Yangon Division). ELSEWHERE: central and
northeastern India throughout all of southeastern Asia including southern China, Thailand,
Cambodia, Laos, Vietnam, and Malaysia, to western Indonesia (Java, Sumatra, Kalimantan).
Hasitat.— In Myanmar, this species has been found primarily in low-lying regions with ele-
vations from close to sea level to around 300 m. Historical records, however, indicate elevations
up to 2300 m (Schleich and Kastle 2002). Most of the individuals encountered have been found in
degraded habitat in the vicinity of villages and agriculture (including paddy). Several have been
found along or near streams. Elsewhere, it is recorded from a diverse array of habitat types (see
Pawar and Birand 2001; Das 2002; Schleich and Kastle 2002). Active at night.
Bungarus flaviceps Reinhardt, 1843
Bungarus flaviceps Reinhardt, 1843:267, pl. iii, fig. 4 (Type locality: Java; Holotype: ZMC R65301).—
Smith, 1943:410.— Toriba, 1993:119.— David and Ineich, 1999:68.
DIAGNOSTIC CHARACTERS.— Dorsal
scales in 13 longitudinal rows at midbody;
expanded neural crest of vertebrae forms dis-
tinct ridge down back and tail; subcaudal
scutes undivided, anteriorly those near the tip
divided; ventrals: o 220— 236, 2 193-217;
subcaudals: 3 47-53, 2 42— 54. Black above;
orange-yellow dorsal stripe often present;
interstitial skin orange-yellow giving appear-
ance of longitudinal stripes; head orange-yel-
low; tail and posterior part of body orange-yel-
low; belly orange or yellow, sometimes edged
with brown. (After Smith, 1943:411.) Total
length 1850 mm; tail length 220 mm.
DISTRIBUTION.— MYANMAR (Mapp.
458): Tanintharyi Division. ELSEWHERE: Thailand, Malayasia, Cambodia, Vietnam, western
Indonesia.
Hasitat.— In Borneo and Thailand this species is found primarily in forested areas from sea
level to around 900 m in elevation. In Sumatra, it is reported to inhabit low lying hills with a pref-
erence for tropical wet forests (David and Vogel 1996). It is generally found under leaf litter and
beneath logs. Active at night (Cox et al 1998; Stuebing and Inger 1999).
REMARKS.— In Myanmar, this widely distributed Malaysian species has been recorded only
from the extreme south, in the vicinity of Myeik (formerly Mergui) and Pyin Mountain.
Bungarus magnimaculatus Wall and Evans, 1901
Bungarus caeruleus magnimaculatus Wall and Evans, 1901:611 (Type locality: Meiktila, Upper Burma;
Holotype: BMNH 1908.6.23.90).
Bungarus magnimaculatus, Smith, 1943:417.— Toriba, 1993:120.— David and Ineich, 1999:69.
DIAGNOSTIC CHARACTERS.— Dorsal scales in 15 longitudinal rows at midbody; subcaudal
scutes undivided throughout; middorsal row of scales (vertebrals) strongly enlarged, as broad as or
broader than long; tail tapering, terminating in a point; ventrals 214~-235; subcaudals 40-48.
Dorsum with 11—14 broad, white crossbars, as wide as the black interspaces, the centers of each of
the scales spotted with black; belly uniformly white. Total length 1300 mm; tail length 150 mm.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 423
DISTRIBUTION.— MYANMAR (Map p. 458):
Magway, Mandalay,and Sagaing Divisions.
Hasitat.— An endemic to Myanmar,
recent herpetological surveys have found this
species primarily in indaing diptocarp forests
(moist deciduous). The type locality of
Meiktila, Upper Burma (referable to Man-
dalay Division), lies in seasonal dry forest;
thus, this species is likely to occur throughout
the central dry zone. Individuals have been
located in disturbed habitats near villages and
in agriculture areas. Active at night.
Bungarus magnimaculatus. Photo by Joseph B. Slowinski.
Bungarus wanghaotingi Pope, 1928
Bungarus multicinctus wanghaotingi Pope, 1928:3 (Type locality: Yuankiang, Yunnan, China; Holotype:
AMNH 35230).— Toriba, 1993:121.— David and Ineich, 1999:69.
Bungarus multicinctus (part) Smith, 1943:416.
DIAGNOSTIC CHARACTERS.— Dorsal
scales in 15 longitudinal rows at midbody; sub-
caudal scutes undivided throughout; middorsal
row of scales (vertebrals) strongly enlarged, as
broad as or broader than long; tail tapering, ter-
minating in a point; ventrals 209-228, subcau-
dals 44-54 (fide Pope 1935:339, Smith 1943:
416). Belly uniformly white; dorsum with
20-31 light crossbands, crossbars narrower
than black crossbars; 7-11 white crossbars on
tail. Total length 1100 mm; tail length 145 mm.
DISTRIBUTION.— MYANMAR (Map _ p.
458): Kachin State, Rakhine State, Sagaing Division, and Yangon area (doubtful). ELSEWHERE:
China (Yunnan).
Hasitat.— This species has been found in a diverse set of habitats, from bamboo stands in
coastal rainforest to streams in indaing forest (moist deciduous), to subtropical forests in Kachin
State. Individuals have been encountered primarily in degraded habitats near villages and along
roads. Wall (1926:564) noted that it has been found at low to mid-range elevations, at least up to
about 900 m. A mating pair was found at night on 10 September in northern Kachin State. Active
at dusk and at night.
REMARKS.— Closely related to but geographically distant from typical B. multicinctus. From
B. multicinctus, it can be distinguished by the lower number of light cross bands on the body and
tail (31-40 on the body and 9-17 on the tail in B. multicinctus, 20-31 and 7—11 respectively in B.
wanghaotingi). The “multicinctus-wanghaotingi’”’ species group requires study; it is likely polytyp-
ic and several closely related though distinct species likely are hidden presently under the names
multicinctus and wanghaotingi.
=i ay e \ SS
Bungarus wanghaotingi. Photo by Hla Tun.
Genus Calliophis Gray, 1834
Calliophis Gray, 1834, pl. 86, fig. 1 (Type species: Calliophis gracilis Gray, 1834).
Maticora Gray, 1834, pl. 86, fig. 2 (Type species: Maticora lineata Gray, 1834 [=Aspis intestinalis Laurent,
1768]).— Slowinski, Boundy, and Lawson, 2001:239.
424 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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REMARKS.— Bourret (1935:414) lists “Burmanie” in his distribution statement for Maticora
bivirgata, but Smith (1943:419) observed, “I do not know of any authentic records of the occur-
rence of this Malayan genus, now known as Maticora, within the area covered by this work.”
Toriba (1993:151—152) also includes Myanmar in his range statement for M. bivirgata (and, as an
aside, he places C. maculiceps in Maticora) but he, too, does not cite new evidence to justify its
inclusion. Under the circumstances, we have chosen to remove M. bivirgata from the faunal list for
Myanmar inasmuch as to date none have shown up in the survey collections that have been con-
ducted during the past four years.
Regarding the status of the nominal genus Maticora, recent work has shown that the genus and
its included species, M. bivirgatus, M. intestinalis, M. maculiceps (fide Toriba 1993), and M.
nigrescens, are properly placed in the genus Calliophis (Slowinski, Boundy, and Lawson 2001).
Calliophis maculiceps Giinther, 1858
Elaps maculiceps Giinther, 1858:232 (Type locality: East Indies; Holotype: BMNH 58.4.20.6).
Callophis maculiceps, Smith, 1943:420.
Maticora maculiceps, Toriba, 1993:153.— David and Ineich, 1999:128.
Calliophis maculiceps, Slowinski, Boundy, and Lawson, 2001:235—241.
DIAGNOSTIC CHARACTERS.— Body scales
in 13 parallel longitudinal rows, not obliquely
disposed; middorsal (vertebral) scales not
enlarged); preocular in contact with nasal; ven-
trals: o 174-186, 2 189-203; subcaudals: 3
25-31, 2 21-25; body above brown to reddish
brown, with black spots, the latter arranged lon-
gitudinally along each side of the back; head
and nape black with some yellow markings [= 8) See os Oe Ae
including a yellow spot on each side of the Calliophis maculiceps. Photo by Hla Tun.
occiput; upper labials yellow; tail, below, pale
blue or gray. Total length 1300 mm; tail length 150 mm.
DISTRIBUTION.— MYANMAR (Map p. 459): as far north as 20°N. ELSEWHERE: Thailand,
Cambodia, Laos, Malaysia.
HABITAT.— Cox et al. (1998) report this species from low elevation forests. The lone individ-
ual encountered during our recent surveys was found at night near a stream in a rubber and
beetlenut plantation at an elevation of 43 m. The surrounding habitat is coastal rainforest and sem1-
evergreen forest. In Thailand, it is usually found under vegetation, rocks or logs (Cox 1991).
Genus Naja Laurenti, 1768
Naja Laurenti, 1768 (Type species: Naja lutescens Laurenti, 1768 [=Coluber naja Linnaeus, 1758}).
Naja kaouthia Lesson, 1831
Naja kaouthia Lesson, 1831:122 (Type locality: Bengal; Holotype: unknown).— Toriba, 1993:187.— David
and Ineich, 1999:159.
Naja naja kaouthia, Smith, 1943:428, 431.
DIAGNOSTIC CHARACTERS.— Body scales smooth, arranged in 19-21 (usually 21) longitudi-
nal rows at mid-body; throat pale, scarcely any dark mottling, often followed by a single dark band,
ventrolateral throat spots distinct; remainder of venter either pale or increasingly cloudy with dark-
er pigmentation towards the rear; in adults, hood markings usually distinct, usually a pale, oval or
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 425
circular marking, with a dark center (see fig.,
left image) and occasionally a narrow dark
outer border; occasionally | or 2 dark spots are
present in the pale oval; fangs not modified for
spitting, venom discharge orifice large; ventrals
164-196; subcaudals 43-58. Total length 1500
mm; tail length 230 mm (according to Smith
[1943:429] larger specimens have been record-
ed, but they are rare).
DISTRIBUTION.— MYANMAR (Map p. 459):
widely distributed throughout the country in
wetter habitats (Ayeyarwady Division, Chin State, Kachin State, Magway Division [part], Rakhine
State, Sagaing Division [part], Yangon Division); in the drier central region, spanning Mandalay,
Magway [part], and Sagaing [part] Divisions, it is replaced by Naja mandalayensis (q.v.).
ELSEWHERE: Nepal, Bangladesh, northeastern India (Assam), Thailand (also in the wetter areas),
northern Malaysia, Cambodia, southern Laos and southern Vietnam, southwestern China (Sichuan,
Yunnan).
Hasitat.— Widely distributed in Myanmar except in the central dry zone (dry and moist
deciduous forests) where its close relative, N. mandalayensis, occurs. Naja kaouthia is often
encountered in villages, in agricultural areas, and grasslands, but it is also met with in primary
coastal rainforest. It has also been encountered swimming in lakes and rivers, as well as beneath
rocks and in the burrows of other animals. Additionally, Cox (1991) observes that it can climb
trees. In Myanmar, it has been recorded from sea level to 820 m. Although most active at dusk and
night, it may be encountered during the daylight hours as well.
Naja kaouthia. Photo by Hla Tun.
Naja mandalayensis Slowinski and Wiister, 2000
Naja naja kaouthia, Smith, 1943:431 (part).
Naja mandalayensis Slowinski and Wiister, 2000:257-270, figs. 1-2 (Type locality: near Monywa [22°13'N,
95°20E], Sagaing Division, Myanmar; Holotype: CAS 207097).
Hood markings in Naja kaouthia (left) and Naja mandalayensis
(right). Photos by Dong Lin.
] Underside of throat and anterior third of body color patterns for
) Naja kaouthia (left) and Naja mandalayensis (right). Photos by
Dong Lin.
426 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DIAGNOSTIC CHARACTERS.— Underside of chin and throat dark, set off from first dark band
by 2 to 4 ventrals that are either pale or at least less densely mottled, followed by 2 or 3 broad dark
bands, the remainder of the venter is pale with occasional dark mottling; hood without markings
(see fig., right image) or, if present, scarcely discernible; fangs modified for spitting, venom dis-
charge orifice small; ventrals: 173-185 (d 173-185, 2 182-185); subcaudals: 50-58 (3 56-58,
50-56). Total length 828 mm; tail length 152 mm (dimensions of holotype, a rather small speci-
men, as recorded by Slowinski and Wiister; larger individuals undoubtedly occur).
DISTRIBUTION.— MYANMAR (Map p. 459): Central dry zone, including parts of Sagaing,
Mandalay, and Magway Divisions.
HABITAT.— This species appears to be restricted to the central dry zone. Most individuals have
been found in the vicinity of villages and agricultural lands.
Genus Ophiophagus Ginther, 1864
Ophiophagus Giinther, 1864 (Type species: Hamadryas elaps Giinther, 1858 [= Naja hannah Cantor, 1836)).
Ophiophagus hannah (Cantor, 1836)
Hamadryas hannah Cantor, 1836:187 (Type locality: Sandarbans, near Calcutta, India; Holotype: Unknown).
Naja hannah, Smith, 1943:436, fig. 140.
Ophiophagus hannah, Toriba, 1993:195.— David and Ineich, 1999:171.
DIAGNOSTIC CHARACTERS.— Body scales smooth, obliquely arranged, in 17—19 rows on the
neck, 15 at midbody; middorsal (vertebral) row and outer 2 lateral rows larger than others; ventrals
240-254; subcaudals 84-104, anterior scutes undivided; juveniles are usually dark brown or black
with white or yellow cross bars, anteriorly the bars are chevron-shaped but straighten out posteri-
orly; with age, the light pattern disappears, and older adults are uniformily brown although some
indication of the light cross bars persists (in Myanmar, the banded pattern persists in adults); tail
dark olive to black. Total length recorded to 5500 mm, but individuals rarely exceed 4250 mm; tail
length approximately 20% of total length.
DISTRIBUTION.— MYANMAR (Map p. 459): widely distributed (encountered by the Survey
team on occasion at localities in both Ayeyarwady and Mandalay Divisions). ELSEWHERE: widely
distributed throughout Southeast Asia and east to the Philippines and western Indonesia.
Hasitat.— In Myanmar, this species has been found in a variety of habitats, in dense forests,
mangrove swamps, open country, and disturbed areas in the Ayeyarwady Delta, to the dry forests
of the central dry zone to coastal rainforest in Tanintharyi. In Assam, Pawar and Birand (2001) con-
firm its presence in primary forest; in Thailand, Cox (1991) states that it is found in both forests
Site Ca
oe
S yo —_—_--. Le he 2
nd ee _ é wt ; Ophiophagus hannah (juvenile). Photo by John Tashjian.
Ophiophagus hannah (adult). Photo by Hla Tun.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 427
and plantations habitats; and in Peninsular Malaysia and Singapore, Lim and Lee (1989) note that
it occurs in foothill jungles, open grasslands, in rural areas, and along jungle streams. David and
Vogel (1996) state that in Sumatra it ranges from sea level to 1800 m.
Genus Sinomicrurus Slowinski, Boundy, and Lawson, 2001
Sinomicrurus Slowinski, Boundy, and Lawson, 2001:239 (Type species: Elaps macclellandii Reinhardt,
1844).
Sinomicrurus macclellandii (Reinhardt, 1844)
Elaps macclellandii Reinhardt, 1844:532 (Type locality: Assam, India; Holotype: Unknown).
Calliophis macclellandi, Smith, 1943:423.
Hemibungarus macclellandi, David and Ineich, 1999:98.
Hemibungarus macclellandi macclellandi, Toriba, 1993:142.
Sinomicrurus macclellandi, Slowinski, Boundy, and Lawson, 2001:239.
Sinomicrurus macclellandi macclellandi, Hallermann et al., 2002:151.
DIAGNOSTIC CHARACTERS.— Vertebral
series of scales not enlarged; body scales in 13
parallel longitudinal rows; | preocular; 2 pos-
toculars; 7 upper labials; temporals 1 + 1, the
anterior shield in contact with 2 upper labials;
color pattern red or brownish above with a
series of narrow black transverse bars (some-
times a narrow, black vertebral stripe with
transverse black bars restricted to sides of body
[Eastern Himalayas] or black transverse bars
reduced to transverse vertebral spots [Assam, =
Upper Myanmar]) (fide Smith 1943:424); head Sinomicrurus macclellandii. Photo by Nikolai Orlov.
black anteriorly reaching back to the level of
the eyes, followed by a broad white band, which is bordered posteriorly by a black nuchal band;
ventrals: ¢ 182-212, 2 208-244; subcaudals, mostly paired, occasionally a few single: 3 28-36,
2 25-33. Total length: ¢ 635 mm, ? 780 mm; tail length: ¢ 70 mm, ? 60 mm.
DISTRIBUTION.— MYANMAR (Map p. 459): Kachin State south to (?) Yangon Division (based
on questionable record). ELSEWHERE: India (Sikkim, Assam), Thailand, southern China, Vietnam.
Hasitat.— The only specimen of this species from recent surveys was found in the subtrop-
ical forests (Northern Triangle) of Kachin State at an elevation of 526 m during the day. Outside of
Myanmar, this species has been recorded from elevations of 350 m to 2000 m (Schleich and Kastle
2002). According to Das (2002), it is generally nocturnal. Cox (1991) reports that in Thailand it is
usually found under loose soil or vegetation in forests up to 1800 m.
Subfamily HYDROPHIINAE
Genus Laticauda Laurenti, 1768
Laticauda Laurenti, 1768:109 (Type species: Laticauda scutata Laurenti, 1768).
Laticauda colubrina (Schneider, 1799)
Hydrus colubrinus Schneider, 1799:238 (Type locality: None specified; Holotype: ZMB 9078).
Laticauda colubrina, Smith, 1926:6; 1943:443.— Toriba, 1993:146.— David and Ineich, 1999:123.
DIAGNOSTIC CHARACTERS.— Ventrals large, one-third to more than half the width of the body;
428 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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nostrils lateral; nasals separated by internasals;
21-25 longitudinal rows of imbricate scales at
midbody; an azygous prefrontal shield usually
present; rostral undivided; ventrals 213-243;
subcaudals: 3 37-47, 2 29-35 (ventral and
subcaudal counts after Smith 1943:443).
Upper lip yellow. Total length: ¢ 875 mm, ¢
1420 mm; tail length: ¢ 130 mm, 2 145 mm.
DISTRIBUTION.— MYANMAR (Map _ pp.
460): coastal waters, tidal rivers, and ashore
especially along rocky coasts. According to LEI EEIS COE JSD (Oy) IDLO ite.
Smith (1943:444), this species is not commonly met in “Indian and Indo-Chinese waters” though
it is not uncommon at Singapore. Minton (1975:26, table 1) suggests that although rare in the Bay
of Bengal, it may not be uncommon along the Myanmar coast and the west coast of the Malayasian
peninsula. ELSEWHERE: coastal waters of Thailand, Malaysia, and western Indonesia as far east as
Polynesia and north along the east Asian coast and Philippines Islands to southern Japan.
HaBITAT.— Individuals of this species were found on a small, uninhabited island approxi-
mately one mile off the Rakhine coast in the Bay of Bengal. They were seen at rest during the day
at low tide in rock crevices. Surrounding waters were rich in large corals. In New Caledonia,
Ineich and Laboute (2002) report that it is often found inshore under vegetation. It has been found
at depths of more than 60 m, but it appears to prefer depths of less then 20 m (Ineich and Laboute
2002). Active day and night (Ineich and Laboute 2002).
Laticauda laticaudata (Linnaeus, 1758)
Coluber laticaudatus Linnaeus, 1758:222 (part) (Type locality: “in Indiis”; Holotype: NHRM 87-88).
Laticauda laticaudata, Smith, 1926:4; 1943:442.— Toriba, 1993:146.— David and Ineich, 1999:124.
DIAGNOSTIC CHARACTERS.— Ventrals large, one- Foo cog i 1
ee aS.
7]
third to more than one half the width of the body; nos-
trils lateral; nasals separated by internasals; 19 longi-
tudinal rows of imbricate scales at midbody; no azy-
gous prefrontal shield; rostral undivided; ventrals
225-243; subcaudals: 3 38-47, 2 30-35 (ventral and
subcaudal counts after Smith 1943:443). Upper lip
dark brown. Total length: ¢ 910 mm, 2 1070 mm; tail
length: ¢ 110mm, 2 110 mm.
DISTRIBUTION.— MYANMAR (Map p. 460):
Rakhine State. Smith (1943:443) states that it is “rare
in the Oriental region (Calcutta and Little Nicobar
Harbour).” On the other hand, Minton (1975:26, table
1) suggests that although rare in the Bay of Bengal, it
may not be uncommon along the Myanmar coast and
the west coast of the Malaysian peninsula.
ELSEWHERE: western Indonesia (Sumatra and Java) to
Australia, Melanesia and Polynesia, and north along
the east coast of Asia to southern Japan.
HABITAT.— This species has been found near the
¥ i. se AN
Laticauda laticaudata. Photo by John Tashjian.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 429
mouth of a small freshwater stream along the Rakhine coast. The coast in this area is awash with
exposed coral reef and mangrove forest. In Taiwan, Mao and Chen (1980) reported that it is often
found near fresh water. Active day and night (Ineich and Laboute 2002).
Genus Disteira Lacépéde, 1804
Disteira Lacépéde, 1804 (Type species: Disteira doliata Lacépéde, 1804 [=Hydrus major Shaw, 1802]).
REMARKS.— At this time, we have chosen not to follow McDowell (1972) and continue to
recognize the nominal genus Enhydrina, which he referred to the genus Disteira, and its included
species, E. schistosa. We do so with some hesitation because we believe that McDowell was like-
ly closer to the truth in synonymizing Enhydrina with Disteira and that the current arrangement
merely perpetuates an unfortunate case of paraphyly. Pending further studies, however, we have
chosen the conservative approach and follow McCarthy (1993) who recognizes the genus
Enhydrina (q.v.) with its included species, E. schistosa and E. zweifeli.
Disteira nigrocincta (Daudin, 1803)
Hydrophis nigrocinctus Daudin, 1803:380 (Type locality:
Sundarbans, Bengal; Holotype: BMNH 1946.1.10.13, but
original description based on Russell, 1801, p. 7, pl. 6).—
Smith, 1926:44, fig. 18; 1943:452.— David and Ineich,
1999-115!
Disteira nigrocincta, McDowell, 1972:239-244.— McCarthy,
1993:226.
DIAGNOSTIC CHARACTERS.— Mental scute large, not
concealed in mental groove; 1—3 maxillary teeth behind
fangs; 7—9 palatine teeth, similar in size to ptergyoid
teeth; head with yellow supraorbital stripe surrounding a
blackish crown patch that extends forward to prefrontals;
olive to brown above with 40 to 60 narrow dark annuli;
yellowish below; 27—33 scale rows around neck, 39-45
around midbody, imbricate and keeled; ventrals 296—330,
distinct throughout but not twice as large as adjacent
scales; preanal scales enlarged. Total length 1080 mm, tail
length 125 mm.
DISTRIBUTION.— MYANMAR: coastal waters and tidal
streams bordering the Bay of Bengal. ELSEWHERE: known
only from the Bay of Bengal and adjacent coasts.
Hasitat.— Little is known about this species. Other
members of this genus are diurnal and are found in deep, Disteira nigrocincta. From Fayrer (1874, pl. 25).
turbid, sandy bottom waters (O’Shea 1996).
Genus Enhydrina Gray, 1849
Enhydrina Gray, 1849:47 (Type species: Enhydrina valakadyen Gray, 1849 [= Hydrus valakadyn F. Boie,
1827 = Hydrophis schistosus Daudin, 1803}).
Remarks: See note above under the genus Disteira.
430 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Enhydrina schistosa (Daudin, 1803)
Hydrophis schistosus Daudin, 1803:386 (Type locality: Tranquebar, South India; Holotype: BMNH
1946.1.10.7, but original description based on Russell, 1801, pl. 10).
Enhydrina schistosa, Smith, 1926:36, fig. 17; 1943:449, fig. 144.— McCarthy, 1993:227.— David and
Ineich, 1999:92.
Disteira schistosa, McDowell, 1972:239-244.
DIAGNOSTIC CHARACTERS.— Mental scute
small, partly concealed within mental groove; 3-4
maxillary teeth behind fangs; 5—6 palatine teeth,
palatine teeth larger than pterygoid teeth; no subor-
bital stripe; young dark gray above, whitish below,
with dark gray or black annuli; pattern disappears
in adults which are uniform gray in color; scales
around body variable, in Bay of Bengal, scales
around neck, males 43-52, females 48-55, scales
around midbody, 3 53-60, 2 55-65, scales some-
what imbricate or juxtaposed, with short central
keel; ventrals 262—322; preanal scales only slightly
enlarged. Total length 1400 mm, tail length 180 Enhydrina schistosa. From Fayrer (1874, pl. 18).
mm (but rarely exceeding 1100 mm total length).
DISTRIBUTION.— MYANMAR (Map p. 460):
coastal waters (two specimens, one in the
Myanmar Biodiversity Museum, the second at the
California Academy of Sciences come from
Ayeyarwady Division coastal waters). ELSEWHERE:
coastal waters from the Persian Gulf east to
Indonesia, New Guinea and northern Australia, and
north from Malaysia to the Philippines.
HaBitTaT.— According to Cogger (1975), in [iggseeeesaaes : z
Australia this species is often found in rivers, but it Enhydrina schistosa. Photo courtesy Indraneil Das.
is not clear if they are found in the brackish waters
of the tidal basins or further away from the estuaries upstream in freshwater. Cox et al. (1998) like-
wise report that in Thailand E. schistosa is sometimes found in estuaries and rivers as well as
coastal waters, but again it is not clear how far upstream they go. According to O’ Shea (1996), this
species prefers water of depths from less than 5 m to a maximum of around 30 m. The individuals
encountered in our recent surveys were caught in nets at the mouth of a river. Active during day
and night (O’Shea 1996).
Genus Hydrophis Latrielle, 1802
Hydrophis Latrielle, 1802:193 (Type species: Hydrus fasciatus Schneider, 1799).— Smith, 1926:40; 1943:
451.— McCarthy, 1993:229.
Hydrophis atriceps Gimnther, 1864
Hydrophis atriceps Giinther, 1864:371, fig. (Type locality: Siam; Syntypes: BMNH 1946.1.2.62,
63.9.29.5).— McCarthy, 1993:230.— David and Ineich, 1999:104.
Hydrophis fasciatus atriceps, Smith, 1926:97, fig. 27; 1943:465.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 43]
DIAGNOSTIC CHARACTERS.— Head small, body long and slender anteriorly; scales on thickest
part of body subquadrangular or hexagonal in shape, juxtaposed or slightly imbricate; 5-6 maxil-
lary teeth behind fangs; 2 anterior temporals; body scales in 25—30 (usually 27-29) rows around
the neck, 39-49 (usually 43-45) around midbody (increase in number of rows from neck to mid-
body 12-21, usually 14-18); ventral scales 323-452 (average 366 or less); anterior part of body
including head and neck dark olive to black with pale oval yellowish spots on sides, sometimes
connected as crossbars; posterior, grayish; below whitish; dark rhomboidal spots may extend down
the sides of the body and form complete annuli in young. Total length ¢ 1100 mm, 2? 990 mm; tail
length ¢ 100 mm, 2 75 mm.
DISTRIBUTION.— MYANMAR (fide Toriba 1993), but according to Smith (1943:465), H. atri-
ceps occurs from the Gulf of Siam eastward and is not known to the west. David and Ineich
(1999:105) do not include Myanmar in its recorded range. All references to H. atriceps from the
Bay of Bengal are most probably H. fasciatus, with which H. atriceps has long been associated.
HasitaT.— Smith (1926) reports this species to be common at the mouths of rivers.
REMARKS.— This species is so similar in appearance to H. flaviceps that the two have been
regarded as conspecific, though treated as distinct subspecies (Smith 1926:97 and 1943:465). The
differences between them are given in the diagnoses. Hydrophis atriceps should be removed from
the Myanmar faunal list.
Hydrophis caerulescens (Shaw, 1802)
Hydrus caerulescens Shaw, 1802:561 (Type locality: Indian Ocean [Vizagapatam {=Visakhapatnam }]; Holo-
type: BMNH 1946.1.3.90).
Aydrophis caerulescens, Smith, 1926:90; 1943:463.— McCarthy, 1993:232.— David and Ineich, 1999:106.
DIAGNOSTIC CHARACTERS.— Scales on thickest part of body quadrangular or hexagonal in
shape, feebly imbricate or juxtaposed; 14—18 maxillary teeth behind front fangs; 2 anterior tempo-
rals; scales in 31-43 rows on the neck, 38—54 around midbody (increase from neck to midbody
6-14); ventrals 253-334, distinct throughout though not twice as large as adjacent body scales;
bluish gray above, whitish below, with 40-60 broad bands, about twice as wide as interspaces,
tapering ventrally (in older adults, bands become indistinct).
DISTRIBUTION.— MYANMAR: coastal waters, especially abundant in the Mergui Archipelago
(Tanintharyi Division). ELSEWHERE: both west and east coasts of India (vicinity of Bombay and
Karwar in the west and from Madras to the mouth of the Ganges on the east coast) east through
Straits of Malacca to the Gulf of Siam to southeastern China and western Indonesia.
HasitTat.— No data available.
Hydrophis cantoris Giinther, 1864
Hydrophis cantoris Giinther, 1864:374, fig. (Type locality: Penang, Malaysia; Holotype: BMNH
1946.1.18.30)— McCarthy, 1993:232.— David and Ineich, 1999:106.
Microcephalophis cantoris, Smith, 1926:124, fig. 35; 1943:475.
DIAGNOSTIC CHARACTERS.— Head small, body long and slender anteriorly; scales on thickest
part of body juxtaposed; 5—6 maxillary teeth behind fangs; 23-25 (rarely 21) scale rows around
neck, 41-48 around thickest part of body (increase from neck to midbody 18-24); ventrals divid-
ed by a longitudinal fissure; prefrontal in contact with third upper labial; ventrals 404468. Total
length ¢ 1450 mm, ? 1880 mm; tail length ¢ 120 mm, ? 140 mm.
DISTRIBUTION.— MYANMAR: coastal waters. ELSEWHERE: coastal waters from Pakistan
(Karachi) east, including India, Sri Lanka, Thailand and Malaysia.
Hasitrat.— No data available.
432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Hydrophis fasciatus (Schneider, 1799)
Hydrus fasciatus Schneider, 1799:240 (Type locality: East Indies; Syntypes: ZMB 2836-2837).
Hydrophis fasciatus, Smith 1926:94; 1943:464.— McCarthy, 1993:234.— David and Ineich, 1999:109.
DIAGNOSTIC CHARACTERS.— Head small, body long
and slender anteriorly; scales on thickest part of body sub-
quadrangular or hexagonal in shape, juxtaposed or slightly
imbricate; 5—6 maxillary teeth behind fangs; 2 anterior tem-
porals; body scales in 28-33 rows around the neck, 47-58
around midbody (increase in number of rows from neck to
midbody 20-27); ventral scales 414-514 (average 460);
anterior part of body including head and neck dark olive to
black with pale oval yellowish spots on sides, sometimes
connected as crossbars; posterior, grayish; below whitish;
dark rhomboidal spots may extend down the sides of the
body and form complete annuli in young. Total length 3
1100 mm, ¢ 990 mm; tail length ¢ 100 mm, 2 75 mm.
DISTRIBUTION.— MYANMAR: coastal waters to the
Straits of Malacca. ELSEWHERE: common along east coast
of India (said to be rare along the west coast but it has been
reported from as far west as Karachi).
HABITAT.— Cox et al (1998) report this nocturnal
species inhabits shallow coastal waters.
REMARKS.— So similar in appearance to H. atriceps
that the two have been regarded as conspecific, though
treated as distinct subspecies (see Smith 1926:97 and
1943:465) (see also above, Remarks, under H. atriceps). Hyd ophis facia Eton EA ee eam
Hydrophis gracilis (Shaw, 1802)
Hydrus gracilis Shaw, 1802:560 (Type locality: Unknown; Holotype: BMNH 1946.1.17.37).
Microcephalophis gracilis, Smith, 1926:121; 1943:472, fig. 150.
Hydrophis gracilis, McCarthy, 1993:234.— David and Ineich, 1999:110.
DIAGNOSTIC CHARACTERS.— Head small, body long and slender anteriorly; scales on thickest
part of body juxtaposed; 5-6 maxillary teeth behind fangs; 17—21 scale rows around neck, 30-36
around thickest part of body (increase from neck to midbody 18-24); ventrals divided by a longi-
tudinal fissure; prefrontal in contact with third upper labial; ventrals 220-287. Total length ¢ 950
mm, 2 1025 mm; tail length ¢ 80 mm, 2 95 mm.
DISTRIBUTION.— MYANMAR: coastal waters. ELSEWHERE: coastal waters from the Persian Gulf
east to India, Sri Lanka, Thailand, Malaysia, Vietnam, China, Taiwan, Indonesia (Sumatra and
Java), Australia, Melanesia.
HABITAT.— Reported to inhabit deep turbid offshore waters (O’Shea 1996).
REMARKS.— Smith (1943:473) provides information on the geographic variation in scale
counts for this species. The values included in the diagnosis above apply only to those populations
inhabiting the coastal waters of Myanmar and along the east coast of India.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 433
Hydrophis obscurus Daudin, 1803
Hydrophis obscura Daudin, 1803:375 (Type locality: Sandbarbans [sic], India; Holotype: BMNH 1946.1.9.27
[but original description based on Russell, 1801, pl. 8])— Smith, 1926:66; 1943:457.— McCarthy,
1993:238.— David and Ineich, 1999:115.
DIAGNOSTIC CHARACTERS.— Scales on
thickest part of body with rounded or bluntly
pointed tips, imbricate throughout; 5—7 maxil-
lary teeth behind fangs; body elongate, ventrals
distinct throughout, 300-338; 6—7 upper labi-
als; 1 anterior temporal; 19—23 scale rows on
neck, 29-37 on thickest part of body (increase
from neck to midbody 8-14); young black with
35-55 bright yellow or whitish dorsal bars,
posteriorly becoming complete bands _ that
encircle body, head with curved yellow mark-
ing extending from snout to either side of pari-
etal scales; adults markings become indistinct and older individuals almost uniform grayish above,
yellowish below. Total length ¢ 1190 mm, 2 1200 mm; tail length o 135 mm, ? 110 mm.
DISTRIBUTION.— MYANMAR: coastal waters especially of the Ayeyarwady Division and south
(Tanintharyi Division). ELSEWHERE: east coast of India.
HasitTaT.— This species is reported to occur mainly in brackish waters (Smith 1926), and his-
torical records indicate that it can be found at the mouths of rivers (ibid. ).
Hydrophis obscurus. From Fayrer (1874, pl. 26).
Hydrophis ornatus (Gray, 1842)
Aturia ornata Gray, 1842:61 (Type locality: Indian Ocean; Holotype: BMNH 1946.1.23.72).
Hydrophis ornatus, Smith, 1926:6.— David and Ineich, 1999:116.
Hydrophis ornatus ornatus, Smith, 1943:460.— McCarthy, 1993:239.
DIAGNOSTIC CHARACTERS.— Scales on thickest part of body more or less hexagonal in shape,
feebly imbricate or juxtaposed; 10-13 maxillary teeth behind fangs; head large; body robust, not
elongate, greatest diameter posteriorly about twice that of the neck; 1 preocular; 2 postoculars; 2
anterior temporals; 7-8 upper labials; scale rows on neck: 3 28-37, 2 31-45, on thickest part of
body, ¢ 33-45, 2 39-55 (increase from neck to midbody 4—12); ventrals distinct throughout, in 3
209-260, in 2 236-312, anteriorly ventrals about twice as large as adjacent scales, narrowing pos-
teriorly; above grayish or light olive to almost white with broad dark bars or rhomboidal spots sep-
Pa r a EEE TE
te tgs
Hydrophis ornatus. Photos by Dong Lin.
434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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arated by narrow interspaces; below yellowish or whitish. Total length ¢ 950 mm, ¢ 860; tail
length ¢ 115 mm, 2 80 mm.
DISTRIBUTION.— MYANMAR (Map p. 460): coastal waters. ELSEWHERE: widely distributed
from the Persian Gulf east to New Guinea and Australia and north along the coast of China to the
Ryukyu Archipelago.
HasitaT.— Reported to inhabit clear waters with coral reefs, as well as turbid rivers and estu-
aries (O’Shea 1996; Ineich and Laboute 2002). Active at night and day (Ineich and Laboute 2002).
REMARKS.— Smith (1943:461) recognized two subspecies, H. 0. ornatus and H. o. ocellatus,
the latter in the coastal waters of Australia. In 1993, McCarthy synonymized H. o. ocellata with H.
o. ornatus but recognized H. 0. maresinensis, described by Mittleman in 1947 to accommodate the
populations found off the coast of China, Taiwan and the RyuKyu Islands.
Hydrophis spiralis (Shaw, 1802)
Hydrus spiralis Shaw, 1802:564 (Type locality: Indian Ocean; Holotype: BMNH 1946.1.6.94).
Hydrophis spiralis, Smith, 1926:48; 1943:453.— McCarthy, 1993:240.— David and Ineich, 1999:118.
DIAGNOSTIC CHARACTERS.— Scales on thickest part of body with rounded or pointed tips,
imbricate; 6-7 maxillary teeth behind fangs; normally 1 anterior temporal; 6—8 upper labials;
25-31 scale rows around neck, 33—38 around midbody (increase from neck to midbody 4-8); ven-
trals 295-362, distinct throughout, about twice as broad as adjacent body scales; yellowish or yel-
lowish-green above, dorsal scales edged with black, 41-46 narrow black bands encircle body, the
bands usually less than one-third the width of the lighter interspaces; head in young black with yel-
low horseshoe-shaped marking, in adult head usually yellow. Total length ¢ 1620mm, 2 1830
mm; tail length ¢ 140 mm, 2 120 mm.
Hydrophis spiralis
From Fayrer (1874, pl. 21).
a
Hydrophis stricticollis
From Fayrer (1874, pl. 28).
a
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 435
DISTRIBUTION.— MYANMAR: coastal waters and tidal rivers. ELSEWHERE: Persian Gulf east to
central Indonesia (Sulawesi) and north to the Philippines.
Hasirat.— Little is known about the natural history of this species; it has been reported in
deep water habitats (Ineich and Laboute 2002).
Hydrophis stricticollis Giinther, 1864
Hydrophis stricticollis Giinther, 1864:376, fig. (Type locality: India; Holotype: BMNH 1946.1.6.90).—
Smith, 1926:73; 1943:459.— McCarthy, 1993:241.— David and Ineich, 1999:119.
DIAGNOSTIC CHARACTERS.— Scales on thickest part of body subquadrangular or hexagonal in
shape, feebly imbricate or juxtaposed; 8—11 maxillary teeth behind fangs; head small, body long
and slender anteriorly, posteriorly 2.5 to 3 times thicker than anteriorly; 1 anterior temporal, rarely
divided; 7—8 upper labials, second in contact with prefrontal, 3-4 border eye; 34-41 scale rows
around neck, 45—55 around midbody; ventrals 374452, distinct thoughout, less than twice as large
as adjacent body scales; grayish to olive above, yellowish below, with 45—65 dark bands, widest
dorsally, disappearing with age; head black or olive, yellow markings on snout and along sides of
head. Total length ¢ 1050 mm, 2 1050 mm; tail length 3 140 mm, ? 90 mm.
DISTRIBUTION.— MYANMAR: coastal waters from Rakhine State south to Gulf of Martaban.
ELSEWHERE: east coast of India from Orissa to Bengal, and Bangladesh.
Hasitat.— Historic records exist for this species in rivers in the Bago Division (Smith 1926);
however, not much is known about its habitat preferences.
Genus Kerilia Gray, 1849
Kerilia Gray, 1849:57 (Type species: Kerilia jerdoni Gray,
1849).— Smith, 1926:31; 1943:446.
Kerilia jerdoni Gray, 1849
Kerilia jerdoni Gray, 1849:57 (Type locality: Madras, India;
Holotype: BMNH III.8.1.a).— Smith, 1926:31, fig. 15; 1943:
447, fig. 143— McCarthy, 1993:242.— David and Ineich,
1999:120.
DIAGNOSTIC CHARACTERS.— Body subcylindrical,
nearly uniform diameter throughout; scales keeled and
imbricate, in 17—23 longitudinal rows, 17 on neck, 21—23
(19-21 for the Bay of Bengal) at midbody; head short; pre-
frontals usually not in contact with upper labials; 6 upper
labials, 3-4 bordering eye; 1 pre- and | postocular; | large
anterior temporal; ventrals 225—253 for the Bay of Bengal
and Gulf of Siam (247-278 further east), small, distinct
throughout, usually entire; olive above, yellowish or white
below, with black dorsal spots of crossbars that form com-
plete bands, especially in young. Total length 1000 mm, tail
length 100 mm.
DISTRIBUTION.— MYANMAR: Tanintharyi Division
(Mergui Archipelago). ELSEWHERE: east coast of India and
Sri Lanka east to the Straits of Malacca, the east coast of Kerlia enone Gromit ayn alee
Malaysia to Borneo.
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Genus Lapemis Gray in Hardwicke and Gray, 1834
Lapemis Gray in Hardwicke and Gray, 1834, vol. 2, pl. 87, fig. 2 (Type species; Lapemis hardwickii Gray).
REMARKS.— McDowell (1972) argued that he could not distinguish the genus Thalassophina
Smith (1926) (type species, Thalassophis viperina Schmidt [1852]) from Lapemis. Although we
believe that McDowell’s views deserve serious consideration, again, as with Enhydrina (q.v.), we
have taken the conservative approach and recognize Thalassophina as a distinct genus.
Lapemis hardwickii Gray in Hardwicke and Gray, 1834
Lapemis hardwickii Gray in Hardwicke and Gray, 1834, vol. 2, pl. 87 (Type locality: Penang, Malaysia;
Holotype: BMNH 1946.1.18.39).— Smith, 1926:108, fig. 32, pl. 1, fig. 3; 1943:468, figs. 148-149.
Lapemis curtis hardwickti, McCarthy, 1993:244.
DIAGNOSTIC CHARACTERS.— Body short,
stout, neck region not less than half as thick at
midbody; head large; scales squarish or hexag-
onal, juxtaposed, outer 34 rows larger than
others, scale rows: & 23-31 around neck,
27-35, around midbody, 3 25-27, 2 33-41;
ventrals small, usually distinct anteriorly, not
so posteriorly, in ¢ 114-186, in 2 141-230;
head shields entire, parietals occasionally
divided; nostrils superior, nasals in contact
with one another; prefrontal usually in contact
with second upper labial; 7-8 upper labials,
3-4 bordering eye; | pre- and 1-2 postoculars;
2, rarely 3, anterior temporals; greenish or yel-
low-olive above, whitish below, 35—SO olive to
dark gray dorsal bars, tapering to a point later-
ally, occasionally encircling body, a narrow dark ventral stripe or broad irregular band occasional-
ly present; adults often lack any pattern and are uniform olive to dark gray; head pale olive to black,
yellow markings on snout present or not. Total length 860 mm, tail length 85 mm.
DISTRIBUTION.— MYANMAR (Map p. 460): coastal waters of the Taninthary1 Division (Mergui
Archipelago). ELSEWHERE: southeast coast of India and the Straits of Malacca, east to Australia and
north to China, Philippines, Taiwan, and Japan.
HABITAT.— Known to be active during both day and night, this species is found to inhabit
coral reefs; it also occurs in estuaries, and tidal zone regions with sandy or muddy bottoms (O’Shea
1996). It is usually found at depths of 6 to 15 m, but it has been encountered in deeper waters
(O'Shea 1996).
REMARKS.— Gritis and Voris (1990) do not recognize L. hardwickii as a distinct species, plac-
ing it in the synonymy of L. curtis. McCarthy (1993) recognizes it as a subspecies of L. curtis,
allowing that the nominate form inhabits coastal waters from the Persian Gulf to the shores of west-
ern India, and L. curtis hardwickii ranges from the coastal waters of Sri Lanka and eastern India
east to New Guinea and Australia and north to the coast of China, the Philippines, and Japan (see
also David and Ineich 1999:121—122). Smith (1926:113, 1943:471) argues that L. curtis ranges
from the Persian Gulf to the west coast of India as far as Sri Lanka but that it is unknown along the
east coast of India. We have chosen to follow Smith’s treatment of the two and recognize L. hard-
wickii as a distinct species.
Lapemis hardwickii. (A preserved specimen; CAS-SU 12434.)
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 437
Genus Pelamis Daudin, 1803
Pelamis Daudin, 1803:361 (Type species: Pelamis bicolor Schneider, 1799 [=Anguis platura Linnaeus,
1766]).
Pelamis platurus (Linnaeus, 1758)
Anguis platura Linnaeus, 1766:391 (Type locality: Unknown; Holotype: Unknown).
Pelamis platurus, Smith, 1926:116, fig. 33; 1943:476— McCarthy, 1993:245.— David and Ineich, 1999:174.
DIAGNOSTIC CHARACTERS.— Body com-
pressed, posteriorly more than twice the diam-
eter of the neck; body scales juxtaposed, sub-
quadrangular in shape, in 49-67 rows around
thickest part of body; ventral scales, 264-406,
very small and, if distinct, divided by a longitu-
dinal groove, but usually indistinguishable
from adjacent body scales; head narrow, snout
elongate, head shields entire, nostrils superior,
nasal shields in contact with one another; pre-
frontal in contact with second upper labial; 1—2
pre- and 2-3 postoculars; 2—3 small anterior
temporals; 7-8 upper labials, 4-5 below eye
but separated from border by subocular; color variable but most often distinctly bicolored, black
above, yellow or brown below, the dorsal and ventral colors sharply demarcated from one anoth-
er; ventrally there may be a series of black spots or bars on the yellow or brown background, or the
yellow may extend dorsally so that there is only a narrow middorsal black stripe, or a series of
black crossbars (see Smith 1943:476-477 for a more complete description of the color pattern vari-
ants). Total length ¢ 720 mm, 2 880 mm; tail length ¢ 80 mm, 2? 90 mm.
DISTRIBUTION.— The most widely distributed of all sea snakes ranging from the east coast of
Africa throughout southern and eastern coastal Asia, as far north as southern Siberia, east through-
out Indonesia to Australia and Tasmania. It is also known from the Gulf of Panama and north to
Baja California in western North America, having arrived there probably during the interstadial,
warm periods of the Pleistocene via a circum-Alaska route following the Japanese and California
currents. Occasional strays have shown up in the Galapagos Archipelago to the south and in the
Hawaiian Islands.
Hasitat.— Although primarily a pelagic species, it has also been found in bays and estuaries.
Active during the day and night (O’Shea 1996).
2 ae MSs. ee
# 2 2% E “a eee
Pelamis platurus. Photo by John Tashjian.
Genus Thalassophina Smith 1926
Thalassophina Smith, 1926:33 (Type species: Thalassophis viperina Schmidt 1852).
REMARKS.— The correct name for this genus has been the subject of controversy for some
years. Most recently, David and Ineich (1999:177) have argued that Praescutata is the proper name
to use. The question arises because of uncertainty of just when the errata notice that accompanies
Wall’s original description of the genus was inserted into his publication, before or after distribu-
tion had begun. If before, then the work was “published” with the errata sheet, which becomes part
of the original publication; if after some copies had been distributed, then David and Ineich’s inter-
pretation is justified. David and Ineich state that Wall’s errata “was obviously written after the main
part of the work, and later inserted in distributed copies.” The question in not in when the errata
438 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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sheet was written or even printed, but when it was published, that is available for distribution. For
the present, we choose to recognize Thalassophina as the valid name for the genus.
Thalassophina viperina (P. Schmidt, 1852)
Thalassophis viperina Schmidt, 1852:79, pl. 3 (Type locality: Java; Holotype: ZMH 404).
Thalassophina viperina, Smith, 1926:33, fig. 16.— McCarthy, 1993:247.
Praescutata viperina, Smith, 1943:448.— David and Ineich, 1999:177.
DIAGNOSTIC CHARACTERS .—
Scales hexagonal, juxtaposed, in
27-34 rows on the neck, 37-50 at
midbody; ventrals 226—274, anterior-
ly about half the width of the body,
narrowing posterior to about twice
the width of the adjacent scales, or
slightly less; head shields entire, nos-
trils superior, nasal shields in contact
with one another; prefrontals longer
than broad, not in contact with upper
labials; 1, rarely 2, pre- and 1—2 post-
oculars; 7-9 upper labials, 3—5 bor-
dering eye (sometimes only 3-4 or
4—5); usually 1 anterior temporal,
occasionally 2 or 3; body color, more
or less bicolored, gray above, white
below, the 2 usually clearly demarked
on the sides, often with 25-35 dark
rhomboidal spots, rarely with dark
bands. Total length ¢ 925 mm, 2 820
mm; tail length ¢ 100 mm, 2 80 mm.
DISTRIBUTION.— MYANMAR:
coastal waters. ELSEWHERE: Persian
Gulf east to Gulf of Siam, southern
China and Borneo.
Thalassophina viperina. From P. Schmidt, 1852, pl. 3.
FAMILY VIPERIDAE
Subfamily AZEMIOPINAE
Genus Azemiops Boulenger, 1888
Azemiops Boulenger, 1888 (Type species: Azemiops feae Boulenger, 1888).
Azemiops feae Boulenger, 1888
Azemiops feae Boulenger, 1888:603, pl. 7 (Type locality: Kakhyen Hills, Burma; Holotype: MSNM 30891).—
Smith, 1943:480, fig. 152.— Toriba, 1993:258.— Zhao and Adler, 1993:274.— David and Ineich,
1999:205— McDiarmid, Campbell, and Touré, 1999:230.— Mallow, Ludwig, and Nilson, 2003:14, pl.
iNet
DIAGNOSTIC CHARACTERS.— No sensory pit between nostril and eye: body cylindrical; head
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 439
flattened, above covered with large, symmetri-
cal shields; nostril large, in single completely
differentiated nasal; loreal shield present,
small; 2 pre- and 2 postoculars; eye with verti-
cally elliptic pupil; scales smooth, in 17 longi-
tudinal rows at midbody; ventrals 180-189;
subcaudals 42-53, mostly paired, occasionally
anterior shields undivided; blackish above,
scales often edged with gray, 14-15 narrow
white or pinkish crossbands, sometimes inter-
rupted middorsally, or alternating with one
another laterally; head yellow with a pair of
dark brown to black stripes of somewhat irreg-
ular width extending from prefrontals to the
black color on the neck. Total length 3 925
mm, 2 820; tail length ¢ 100 mm, 2 80 mm.
DISTRIBUTION.— MYANMAR: Kachin
State. ELSEWHERE: southern and central China
(western Yunnan and Shaanxi east to Zhejiang
and south to Guangxi), and northern Vietnam.
Hasirat.— Although little is known about
the natural history of this species, Kardong
(1986) (summarizing Zhao and Zhao 1981)
reports that it inhabits mountainous terrain at é 4
elevations between 1000-2000 m, but it has = eupioi euae TORRE:
also been found in degraded habitats such as
paddy, grassy fields and in and about villages. Historical records of this species in Myanmar indi-
cate that it occurs in the Northern Triangle subtropical forest and the Nujiang Langcang Gorge’s
alpine-conifer and mixed-deciduous forest. For additional comments see Mallow et al. (2003).
Subfamily VIPERINAE
Genus Daboia Gray, 1842
Daboia Gray, 1842:69 (Type species: Daboia elegans Gray, 1842 [=Vipera daboia Daudin, 1803=Coluber
russelit Shaw and Nodder, 1797], fide Smith, 1943:482).
Daboia russelii (Shaw and Nodder, 1797)
Coluber russelii Shaw and Nodder, 1797:291 (Type locality: India; Holotype: BMNH II.I.Ia).
Vipera russelli siamensis, Smith, 1917:223; 1943:484, fig. 153.
Daboia russelli siamensis, Toriba, 1993:268.
Vipera russellii, Zhao and Adler, 1993:278.
Daboia russelii, David and Ineich, 1999:312.— McDiarmid, Campbell, and Touré, 1999:371.— Mallow,
Ludwig, and Nilson, 2003:150, pl. 7.2.
DIAGNOSTIC CHARACTERS.— No sensory pit between nostril and eye; head very distinct from
neck, above covered by small, keeled, imbricate scales, 6-9 between narrow supraoculars; nostril
large, in large nasal shield which, below, is fused to the rostral; eye, with vertically elliptic pupil,
surrounded by 10-15 small scales, 3-4 rows of small scales separating the circumocular scales
from the upper labials; temporals small; 10—12 upper labials; 27—33 longitudinal rows of scales at
440 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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midbody, all except outmost row _ strongly
keeled; ventrals 153-180; subcaudals 41-64,
all paired; color above light brown with 3 lon-
gitudinal series of large black-margined brown
spots or blotches, the vertebral series often
merging to form a chain-like longitudinal
stripe, occasionally an additional longitudinal
series of small dark spots between vertebral
and lateral series; yellowish white below occa-
sionally with dark brown markings. Total ee
length to 1600 mm are not uncommon (fide Daboia russelii. Photo by Hla Tun.
Smith 1943:484).
DISTRIBUTION.— MYANMAR (Map p. 461): Ayeyarwady Division, Bago Division, Magway
Division, Mandalay Division, Sagaing Division, Shan State, Yangon Division. ELSEWHERE: south-
ern China, Taiwan, India, Pakistan, Sri Lanka, Bangladesh, Thailand, Indonesia (Java east to
Lomblen Island). Not reported from Malaysia or Sumatra.
HABitaT.— Primarily inhabits lowland areas. It is common throughout the central dry zone
and the Ayeyarwady delta and is often encountered in agricultural areas and paddies as well as open
grasslands. Active at night.
REMARKS.— Possibly the most common of the dangerously venomous snakes occurring in
southern Asia, and responsible for more than half of all reported snakebites. Several color pattern
variants have been recognized as distinct subspecies: Daboia russelii siamensis from southern
China, central and southern Myanmar, and central Thailand; D. 7 formosensis from eastern China
and Taiwan; and the nominate form from India, Pakistan, and Bangladesh. Its unusual distribution,
especially its erratic distribution in Indonesia, suggests it has been transported in the course of com-
mercial exchanges, likely during the 18th and 19th centuries. Daboia russelii is a prolific breeder
and young could easily have been transported among plants and other products that were frequent-
ly carried about during the early days of colonial expansion.
Arguments over the correct spelling of the species name, i.e., russelii versus russellii, abound
to this day (most recently, see Adler et al. 2000:83, David and Ineich 1999:313, and McDiarmid,
Campbell, and Touré 1999:370). The species was named for Patrick Russell, who spelt his name
with a double “Il.” However, there is no indication in the original publication by Shaw and Nodder,
or any notice subsequently issued, that their use of the single “I” was a lapsus of any sort. Thus, we
follow the original orthography despite the fact that the name of the person being honored is mis-
spelled.
For extended comments on habitat, behavior, reproduction, bite and venoms, see Mallow et al.
(2003).
Subfamily CROTALINAE
Genus Ovophis Burger, 1981
Ovophis Burger in Hoge and Romano-Hoge, 1981:246 (Type species: Trimeresurus monticola Giinther 1864).
Ovophis monticola (Giinther, 1864)
Trimeresurus monticola Giinther, 1864:388 (Type locality: Nepal and Sikkim; Syntypes: BMNH 1946.1.18.76
and 1946.1.19.91.— Pope, 1935:127, pl. 24D—-E and 27A.— Smith, 1943:506, fig. 161.
Trimeresurus monticola monticola, Zhao and Adler, 1993:276.
Ovophis monticola monticola, Toriba, 1993:81.— Hallermann et al., 2002:152.
Ovophis monticola, McDiarmid, Campbell, and Touré, 1999:316.— David and Ineich, 1999:268.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 44]
DIAGNOSTIC CHARACTERS.— Body stout;
snout short, a little more than twice the length
of the diameter of the eye; head covered above
by small scales rather than large shields, scales
usually smooth, feebly imbricate; first upper
labial not fused to nasal, completely separated
by a suture; body scales, smooth or weakly
keeled, in 23—25, occasionally 19 or 21 longi-
tudinal rows at midbody; supraoculars large,
5-9 scales in a line between them; internasals
usually not in contact with one another, sepa-
rated by 2 small suprapostrostral scales; 7—10
upper labials, second usually fused to the scale
bordering the facial sensory pit anteriorly,
fourth and fifth beneath eye but separated from
orbit by 24 series of small scales; ventrals and
subcaudals (Myanmar, northeastern India and
adjacent areas of China and Thailand) 137-176 Ovophis monticola. From Fayrer (1874, pl. 15).
and 36-62 respectively, subcaudals mixed
paired and single, occasionally all unpaired (ventrals and subcaudals for southern China, Vietnam,
Laos: 127—144 and 36—54, and Malaysian Peninsula: 133-137 and 22—28 respectively [fide Smith
1943:509]). Total length ¢ 490 mm, 2 1100; tail length ¢ 80 mm, 2 150 mm.
DISTRIBUTION.— Widely distributed from the eastern Himalayas, Myanmar (Map p. 461),
southeastern Xizang (Tibet) and Yunnan, Thailand, southern China and Taiwan, to Vietnam, the
Malaysian Peninsula to western Indonesia (Sumatra).
Hasitat.— Found in the temperate and subtropical forests of northern Kachin State, from ele-
vations around 1000 m. The altitudinal distribution throughout its range is reported between 700-
2400 m (Schleich and Kastle 2002). Individuals have been found in leaf litter and shifting cultiva-
tion. Pope (1935) reports that this species is common around villages. Crepuscular (Schleich and
Kastle 2002), also secretive and sluggish and most often found tucked away in piles of wood, logs,
and rocks, also in rock crevices (David and Vogel 1966).
REMARKS.— Zhao and Adler (1993:276) and Toriba (1993:82) recognize several subspecies,
the westernmost populations from Nepal, northestern India, Myanmar, and Yunnan and Szechwan
Provinces of China as Ovophis monticola monticola (genus Trimeresurus in Zhao and Adler); O.
m. convictus from Cambodia, Vietnam, Thailand, West Malaysia and western Indonesia; O. m.
Ovophis monticola (JBS 11879) from Kachin, Myanmar. Photo by Hla Tun.
442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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makazayazaya from eastern China and Taiwan; and O. m. zayuensis from the type locality at Zayii
Co, Xizang (Tibet), China. At this time, we take no position on the status of these taxa save that all
are clearly members of a “monticola” species group.
Genus Protobothrops Hoge and Romano-Hoge, 1983
Protobothrops Hoge and Romano-Hoge, 1983:87 (Type species: Trimeresurus flavoviridis Hallowell, 1861).
REMARKS.— In 1983, Hoge and Romano-Hoge described the new genus Protobothrops to
accommodate two species previously placed in the genus Trimeresurus, Trimeresurus jerdonii and
T; mucrosquamatus. Since that time, few authors have followed their recommendation. More
recently, however, Kraus et al. (1996) indicated that preliminary DNA studies support its recogni-
tion, though, as pointed out by McDiarmid et al. (1999:329), it was done with some reservation.
McDiarmid et al. (loc. cit.), thus, chose to take a more conservative approach, pending further stud-
ies, and treated Protobothrops as a synonym of Trimeresurus. In the same year, David and Ineich
(1999:274), also citing Kraus et al., as well as additional but unpublished data (received from N.
Vidal), decided to recognize Protobothrops as a valid genus, though they did note that “the limits
of this genus remain provisional.” Of the eight trimeresurid species David and Ineich refer to
Protobothrops, three occur in Myanmar, P. jerdonii, P. kaulbacki, and P. mucrosquamatus. In this
report, we have chosen to follow David and Ineich’s classification scheme.
Protobothrops jerdonii (Giinther, 1875)
Trimeresurus jerdonii Giinther, 1875:233, pl. 34 (Type locality: Khasi Hills, India; Syntypes: BMNH
196.1.18.66—-68).— Pope, 1935:409, pl. 25.— Smith, 1943:510, fig. 162.— Toriba, 1993:100.— David
and Tong, 1997:26, 28.— McDiarmid, Campbell, and Touré, 1999:336.
Protobothrops mucrosquamatus, Hoge and Romano-Hoge, 1983:86.— David and Ineich, 1999:275.— David,
Captain and Bhatt, 2001:224
DIAGNOSTIC CHARACTERS.— Scales in 21 longitudinal rows at midbody (rarely 23); snout
length a little more than twice diameter of eye; head above, except for large internasals and
supraoculars, covered by small, unequal, smooth scales that are feebly imbricate or juxtaposed;
first labial completely separated from nasal by a suture; internasals separated by 1—2 small scales;
6-9 small scales in line between supraoculars; 7-8 upper labials, third and fourth beneath eye, in
contact with subocular or separated by at most a single series of small scales; ventrals (see Remarks
below): 3 164-188, 2 167-193; subcaudals: 3 50-78, 2 44-76. Total length ¢ 835 mm, 2 990
mm; tail length ¢ 140 mm, 2 160 mm.
DISTRIBUTION.— MYANMAR (Map p. 461): Chin State (Chin Hills), Kachin State. ELSEWHERE:
Protobothrops jerdoni. Variation in color pattern (CAS 215115 [left] and CAS 215015 [right]).
Both individuals from Yunnan Province, China. Photos by Dong Lin.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 443
India (Assam), China (Yunnan, Szechwan, Hupeh).
Hasitat.— This species has been recorded in Myanmar and adjacent areas in Yunnan, from
elevations of 1442 m to just under 2300 m. In Nepal it has been recorded as high as 2800 m
(Schleich and Kastle 2002). Historical records from the Chin Hills and Kachin State as well as
recent records from Kachin State place this species in montane (Chin Hills-Rakhine Yoma) and
temperate (Northern Triangle) forests. Individuals have been found in shifting cultivation. Orlov et
al. (2001) report this species (albeit a different subspecies) as commonly found along rocky streams
in trees, shrubs, and under rocks.
REMARKS.— Both Pope (1935:409) and Smith (1943:510) took note of the distribution of ven-
tral and subcaudal counts among their samples in relation to their geographic origins. Smith, in par-
ticular, notes the following: “Burma, Yunnan (17 examples): V. 3 164-173, 2 167-189; C. 2
50-55 (69), 2 44-61. Burma-Tibet border (12 examples): V. ¢ 181-188, 2 184-193; C. 3 67-78,
2 64-76, paired.” Elsewhere in its range, Maslin (1942) chose to recognize the populations from
eastern and southeastern China as a distinct subspecies, Trimeresurus jerdonii xanthomelas
Giinther, and Klemmer (1963) referred the populations from Vietnam and Cambodia to
Trimeresurus jerdonii bourreti. Toriba (1993:100) recognizes both and assigns the remaining pop-
ulations from Myanmar, northeastern India and the Yunnan and Xizang region of China to the nom-
inate subspecies. The interesting distribution of ventral and subcaudal counts recorded by Smith,
apart from the clear indication of sexual dimorphism, suggests the Myanmar-India-southwestern
Chinese populations deserve further careful study.
Protobothrops kaulbacki (M.A. Smith, 1940)
Trimeresurus kaulbacki Smith, 1940:485, pl. 8, fig. 5 (Type locality: Pangnamdim, north of the Triangle,
Upper Burma; Holotype: BMNH 1946.1.19.23-24); 1943:512.— Toriba, 1993:101— McDiarmid,
Campbell, and Touré, 1999:337.
Protobothrops kaulbacki, Kraus, Mink, and Brown, 1996:769.— David and Ineich, 1999:276.
DIAGNOSTIC CHARACTERS.— Scales in 23—25 longitudinal rows at midbody; body elongate,
head long, massive, with narrow snout; canthus rostralis sharp; single large, squarish loreal; first
upper labial completely separated from nasal by a suture; second upper labial anteriorly bordering
facial pit; 8-10 scales in line between supraoculars; supraoculars usually single, flat, without a ver-
tical projection; dull grayish or olive green with large, blackish rhombohedral dorsal blotches,
either distinct or united to one another; smaller spots on sides; symmetrical yellow lines on head;
subcaudals paired; ventrals 201-212; subcaudals 66-78, some of the anterior scutes may be single,
others paired; hemipenes with spines. Total length ¢ 1340 mm, ? 1410 mm; tail length 3 225 mm,
2 230 mm.
DISTRIBUTION.— MYANMAR (Map p. 461): Kachin State (Pangnamdim).
Protobothrops kaulbacki. Photos by Hla Tun.
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HABITAT.— Little is known of the habitat preferences of this rare snake. The lone individual
of this species from recent work was found at the type locality (Pangnamdin) at an elevation of
1015 m. This region falls at the transition zone between temperate Northern Triangle forests and
eastern alpine shrub and meadow.
Protobothrops mucrosquamatus (Cantor, 1839)
Trigonocephalus mucrosquamatus Cantor, 1839:32 (Type locality: Naga Hills, Assam, India; Holotype:
Unknown; original description probably based on colored drawing [no. 18] in Bodleian Library, Oxford).
Trimeresurus mucrosquamatus, Swinhoe, 1870:411, pl. 31.— Pope, 1935:416, pl. 26.— Smith, 1943:507.—
Toriba, 1993:102.— McDiarmid, Campbell, and Touré, 1999:339.
Protobothrops mucrosquamatus, Hoge and Romano-Hoge, 1983:86.— David and Ineich, 1999:276.
DIAGNOSTIC CHARACTERS.— Scales in 25
longitudinal rows at midbody; scales on upper
surface of head, small, each scale keeled poste-
riorly; internasals 5—10 times size of adjacent
scales, separated by 3-4 scales; supraoculars,
long, narrow, undivided, 14—16 small interocu-
lar scales in line between them; 2 scales on line
between upper preocular and nasal; 9—11 upper
labials, first upper labial separated from nasal
by suture; 2—3 small scales between upper labi-
als and subocular; 2—3 rows of temporal scales
above upper labials smooth, above those scales
keeled; ventrals 200-218; subcaudals 76-91,
all paired; grayish or olive brown above, with -
dorsal series of large brown, black-edged spots UDI IES CONE DALLAS VALGUS lohy IeLe WD.
or blotches, and a lateral series of smaller spots; head above brownish, below whitish; belly whitish
but heavily powdered with light brown; tail brownish (possibly pink in life [fide Smith 1943:507]),
with series of dark dorsal spots; hemipenes spinose. Total length ¢ 1122 mm, 2 1160; tail length
6 195mm; 2 205 mm.
DISTRIBUTION.— MYANMAR (Map p. 461): Kachin State ELSEWHERE: northeastern India and
Bangladesh, China, Taiwan and northern Vietnam.
HABITAT.— Found in the Northern Triangle temperate and subtropical forests of Kachin State
at elevations of 250-1088 m. Individuals have been found near streams either under rocks or in the
leaf litter. Pope (1935) reports that the species is common in hilly and mountainous areas. In
Vietnam, Orlov et al. (2001) state that the species is common around villages and disturbed habi-
tat. Active at night.
Genus Trimeresurus Lacépéde, 1804
Trimeresurus Lacépéde, 1804:209 (Type species: Vipera viridis Daudin, 1803 [=Coluber gramineus Shaw,
1802]).
REMARKS.— See remarks under Genus Protobothrops.
Trimeresurus albolabris Gray, 1842
Trimeresurus albolabris Gray, 1842:48 (Type locality: China; Holotype: BMNH 1946.1.19.85).— Smith,
1943:523, fig. 166 [map].— David and Ineich, 1999:280.— McDiarmid, Campbell, and Touré, 1999:329.
Trimeresurus albolabris albolabris, Toriba, 1993:95.— David and Tong, 1997:25-26.
DIAGNOSTIC CHARACTERS.— Scales in 21 (rarely 19) longitudinal rows at midbody;
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 445
10-11(12) upper labials, the first partially or
completely fused to the nasal; head scales
small, subequal, feebly imbricate, smooth or
weakly keeled; supraoculars narrow, occasion-
ally enlarged, undivided, 8-12 interocular
scales between them; temporal scales smooth;
green above, side of head below eyes yellow,
white or pale green, much lighter than rest of
head; below, green, yellowish or white below, a Trimeresurus albolabris. Photo by Hla Tun.
light ventrolateral stripe present in all males,
absent in females; end of tail not mottled brown; ventrals: 3 155-166, 2 152—176; subcaudals: 3
60-72, 2 49-66, paired; hemipenes without spines. Total length ¢ 600 mm, 2 810; tail length 3
120 mm, 2 130 mm.
DISTRIBUTION.— MYANMAR (Map p. 462): north of 13°N. ELSEWHERE: India (Assam),
Thailand, Cambodia, Laos, Vietnam, China, Malaysia, western Indonesia (as far east as Sulawesi).
HasiraT.— Often found in heavily degraded forest, or in agricultural areas. Documented to
occur in moist mixed deciduous, and subtropical forests (both Northern Triangle and Northern
Indochina) as well as temperate forests. This species is commonly encountered in bamboo stands,
although individuals have also been found on trees, in bushes, on the ground in open fields, and
crossing roads. Elevations range from 60—751 m in Myanmar, although it is recorded as high as
3050 m in Nepal (Schleich and Kastle 2002). Active at night. David and Vogel (1996) note that it
prefers lowland habitats, both forested and open, and when in bushes or trees, usually within 2 or
3 m of the ground.
REMARKS.— Two subspecies have been described, 7. a. insularis Kramer (1977) from eastern
Indonesia (Soe and Timor) and 7! a. septentrionalis Kramer (1977) from Nepal and northwestern
India (Simla).
Trimeresurus erythrurus (Cantor, 1839)
Trigonodactylus erythrurus Cantor, 1839:31 (Type locality: Ganges Delta; Holotype: BMNH 1946.1.19.99).
Trimeresurus erythrurus, Smith, 1943:386, fig. 165.— Toriba, 1993:97.— David and Ineich, 1999:283.—
McDiarmid, Campbell, and Touré, 1999:331.
DIAGNOSTIC CHARACTERS.— Scales in
23-25 longitudinal rows at midbody; first upper
labial partially or completely fused to nasal;
9-13 upper labials, 1-2 rows of scales separate
upper labials from subocular; 11—14 scales in a
line between supraoculars; supraoculars rarely
divided; temporal scales small, strongly keeled;
ventrals: males 153-174, females: 151-180;
subcaudals: ¢ 62-79, 2 49-61, usually paired,
occasionally unpaired shields present among
paired series; head uniform green, dorsum
bright green, light ventrolateral stripe present in Trimeresurus erythrurus. Photo by Dong Lin.
males, present or absent in females (Maslin
[1942:23] says that the ventrolateral stripe is absent, but Smith [1943:524] states that it is present
in males and variable in females), tail spotted with brown; hemipenes without spines. Total length
3 575 mm, 2 1045; tail length ¢ 120 mm, ? 165 mm.
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DISTRIBUTION.— MYANMAR (Map p. 462): Sagaing State, Kachin State, Rakhine State,
Yangon Division. 7: erythrurus is found west of Long. 98°. ELSEWHERE: India (northeastern India
from Bengal to Assam, east of Long. 88°) east through Bangladesh.
Hasirat.— Individuals of this species have been found in rainforests (Myanmar coastal,
Mizarom-Manipur-Kachin) and moist deciduous forest. Elevations are recorded under 200 m.
Individuals have been met with in trees and on the ground near streams. Active at night.
Trimeresurus medoensis Zhao, 1977
Trimeresurus medoensis Zhao in Zhao and Jiang, 1977:66, pl. 2, fig. 9, 1-5 (Type locality: near Ani Bridge,
Motuo, Xizang, at 1200 m; Holotype: CIB 73-II-5208).— Toriba, 1993:103.— McDiarmid, Campbell,
and Touré, 1999:339.— David and Ineich, 1999:287.— David et al., 2001:218.— David, Captain and
Bhatt, 2002:210—226, figs. 1-7.
DIAGNOSTIC CHARACTERS.— Scales
in 17 longitudinal rows at midbody, dorsal
rows 7-11 slightly keeled; 8 upper labials,
first upper labials separated from nasals by
a distinct suture; green or bluish green
above, yellowish white below, the two
separated by a bright bicolored red
(below) and white (above) ventrolateral
stripe (in both males and females), which
occupies the whole of the outermost scale
row and a portion of the second row; ven-
trals less than 150; hemipenes short, thick,
spinose. Total length ¢ 671 mm, ¢ 650; Trimeresurus medoensis. Photo courtesy Ashok Captain.
tail length ¢ 125 mm, 2 115 mm.
DISTRIBUTION.— MYANMAR (Map p. 462): Kachin State (Naung Mon, Rainbow Village,
Myitkina; Alangdunhku). ELSEWHERE: China (Xizang Autonomous Region); India (Arunachal
Pradesh) (after David et al. 2001:218; David et al. 2002:210 ff.).
HABITAT.— The only records of this species in Myanmar are from Kachin State. Three speci-
mens from Nam Ti Valley are referenced in the original description (Zhao 1977), and their identi-
fications are confirmed by David et al. (2001). A specimen from the Mytkina area is also referable
to this species (David et al 2001). David et al. (2001) and Das (2002) report this snake as frequent-
ing bamboo stands in otherwise evergreen forest. The lone specimen from recent collections made
by members of the Myanmar survey team was found near a trail in low elevation evergreen forest
that was interspersed with bamboo stands where the forest had been altered. Active at night (Das
2002).
Trimeresurus popeiorum M.A. Smith, 1937
Trimeresurus popeiorum Smith, 1937:730 (Type locality: Sikkim, India; Holotype: BMNH 72.4.17.137.—
David and Ineich, 1999:288.— David, Captain, and Bhatt, 2002:218, 223.
Trimeresurus popeorum, Smith, 1943:518.—Toriba, 1993:103.— McDiarmid, Campbell, and Touré,
1999:340.
Trimeresurus popeiorum popeiorum, David et al., 1997:27.
DIAGNOSTIC CHARACTERS.— Scales in 21 (rarely 23) longitudinal rows at midbody; 9-11
upper labials, first upper labials separated from nasals by a distinct suture; a single supraocular;
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 447
above green, below pale green to whitish, the
two separated by a bright bicolored orange or
brown (below) and white (above) (males) or
white (females) ventrolateral stripe, which
occupies the whole of the outermost scale row
and a portion of the second row; ventrals
155-169; subcaudals 52—76, in males the base
of the tail enlarged to the level of subcaudals
20-25; hemipenes long and slender, smooth,
without spines. Total length 770 mm, tail
length 170 mm.
DISTRIBUTION.— MYANMAR (Map p. 462):
Bago Division, Mon State, Tanintharyi
Division, and (?)Chin State. ELSEWHERE: Laos; Vietnam; Cambodia; Malaysia; Singapore;
Indonesia (Kalimantan, Sumatra), (?) Thailand (see David et al. [2001:218]).
Hasitat.— In Peninsular Malaysia and Singapore, this nocturnal species is reported from
montane forests (Lim and Lee 1986) with elevations from 900-1500 m (Cox et al. 1998).
Elsewhere, it has been found in hilly areas with wet forest (David and Vogel 1996). In Myanmar,
it has been found in low elevation (less than 500 m) montane and coastal rainforest. Individuals
have been collected on tree limbs overhanging streams and in bushes. Active at night.
REMARKS.— Most often confused with 7. stejnegeri (q.v.), the two have quite distinct
hemipenes, which does not make identification of individuals in the field or in the laboratory any
easier without recourse to (a) male individuals and (b) an examination of the hemipenes. However,
the two species are not known to have overlapping distributions, at least based on available mate-
rials. Also, closely allied to 7. popeiorum is T. yunnanensis (q.v.); ordinarily, the two are more eas-
ily be told apart by the number of midbody scale rows, 21 in T? popeiorum, 19 in T: yunnanensis.
The species name “popeiorum” has been variously spelt “popeiorum” and “popeorum.” In the
original description, it was given as “popeiorum,” but in 1943, in the footnote on p. 518, Smith
states “Popeiorum as originally spelt is a clerical error.” Although a clerical error, yet the use of
“popeorum contravenes Article 32(c)(ii), which states that an original spelling is an ‘incorrect
original spelling’ if “there is in the original publication itself, without recourse to any external
source of information, clear evidence of an inadvertent error... (. . . and use of an inappropriate
connecting vowel are not to be considered inadvertent errors).’”’ Because there is no such evidence
in the original publication, Trimeresurus popeiorum must stand as the appropriate spelling for the
nominal species meant to honor Clifford H. Pope.
Trimeresurus popeiorum. Photo by John Tashjian.
Trimeresurus purpureomaculatus Gray, 1830
Trigonocephalus purpureomaculatus Gray in Gray and Hardwicke, 1830:pl. 81 (Type locality: Singapore;
Holotype: BMNH 1946.1.19.54 and Hardwicke’s sketch no. 158).
Trimeresurus purpureomaculatus purpureomaculatus, M.A. Smith, 1943:520.— Toriba, 1993:104.
Trimeresurus purpureomaculatus, David and Ineich, 1999:280.— McDiarmid, Campbell, and Touré,
1999:42.
DIAGNOSTIC CHARACTERS.— Scales in 25—27 longitudinal rows at midbody; 11—13 upper
labials, the first partially or completely united with the nasal; supraocular very narrow, sometimes
broken into small scales, 12-15 scales between them; head scales small, subequal, tuberculate or
granular; temporal scales keeled; body color highly variable, above olive, grayish, to dark purplish
brown, below whitish, greenish or brown, uniform or spotted with brown; a light line on scale row
448 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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one bordering ventrals present or absent; head
olive, heavily suffused with brown; ventrals: 3
160-179, 2 168-183; subcaudals: 3 74-76, 2
56-63, paired; hemipenes without spines. Total
length ¢ 665 mm, ¢ 900; tail length 3d 125
mm, 2 140 mm.
DISTRIBUTION.— MYANMAR (Map _p.
462): Ayeyarwady Division, Mon and Karen
States south of Lat. 17°N, Tanintharyi
Division. ELSEWHERE: Malaysia, Singapore,
western Indonesia (Sumatra).
Hasitat.— This species has been found
in abundance in the remaining mangrove Trimeresurus purpureomaculatus. Photo by Dong Lin.
forests of the Ayeyarwady Delta. Lim and Lee
(1986) similarly report this species from mangrove forests on offshore islands and mainland penin-
sular Malaysia and Singapore. David and Vogel (1996) report that in Sumatra it is found in man-
grove and coastal swamp habitats, in marshes, and along canals and river banks. Individuals have
been found in trees and shrubs. Crepuscular.
REMARKS.— A population of Asian pit vipers that is closely allied to 7: purpureomaculatus
occurs on the Andaman and Nicobar Islands. In 1868, Theobald described it as a new species, 7!
andersoni. Since then it has been variously treated as a synonym of 7) purpureomaculatus or as a
subspecies of the latter. In view of its complete isolation from mainland T? purpureomaculatus and
its distinguishing features (see Smith 1943:520-—521), we take the position that it is best treated as
a distinct species. We do recognize that it was derived from a population of 7. purpureomaculatus
that inhabits a neighboring area, but which of the neighboring areas we do not know.
Trimeresurus stejnegeri K.P. Schmidt, 1925
Trimeresurus stejnegeri K.P. Schmidt, 1925:4 (Type locality: Shaowu, Fukien, China; Holotype: AMNH
21054).— Pope, 1935:409, pl. 25.— Smith, 1943:517.— McDiarmid, Campbell, and Touré, 1999:344.—
David and Ineich, 1999:290.— David, Vidal and Pauwels, 2001:205, 218, 221.— David, Captain and
Bhatt, 2002:218, 221, 223. — Hallermann et al., 2002:152.
DIAGNOSTIC CHARACTERS.— Scales in 21 longitudinal rows at midbody; 9-11 upper labials,
first upper labials separated from nasals by a distinct suture; a single narrow supraocular, some-
times divided by transverse suture; 11—16 scales in a line between supraoculars; above bright to
dark green, below pale green to whitish, the two separated by a bright bicolored orange or brown
(below) and white (above) (males) or bicolored or white only (females) ventrolateral stripe, which
occupies the whole of the outermost scale row and a portion of the second row; ventrals 150-174,
subcaudals 54-77 (but see David et al. [2002:218 and 2001:215] for geographic distribution of
these data), all paired; hemipenes short, spinose beyond bifurcation. Total length 750 mm, tail
length 145 mm.
DISTRIBUTION.— All records of this species from MYANMAR have been referred to
Trimeresurus yunnanensis (see below). ELSEWHERE: China (central and southeastern), Taiwan,
Vietnam. For possible northeast Thailand occurrence, see David et al. (2001:218-—219).
HABITAT.— Pope (1935) reports that this nocturnal species is very common around fast-flow-
ing streams. Elevations are recorded from 500-900 m (ibid.).
REMARKS.— See remarks under 7: yunnanensis (below).
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 449
Trimeresurus yunnanensis K.P. Schmidt, 1925
Trimeresurus yunnanensis K.P. Schmidt, 1925:4 (Type locality: Tengyueh [=Tengchiong Co.], Yunnan, China;
Holotype: AMNH 21058).— David and Tong, 1997:26—27.— David and Ineich, 1999:294.— David,
Vidal and Pauwels, 2001:218.— David, Captain and Bhatt, 2002:218, 221, 223.
Trimeresurus stejnegeri, Smith, 1943:517 (part).
Trimeresurus stejnegeri yunnanensis, Maslin, 1942:22.— Toriba, 1993:106.
DIAGNOSTIC CHARACTERS.— Scales in 19 (rarely 21) longitudinal rows at midbody and
19(—21) on neck; 9-10 (rarely 11) upper labials, first upper labials separated from nasals by a dis-
tinct suture; a single narrow supraocular, sometimes divided by transverse suture; 11—16 scales in
a line between supraoculars; above bright to dark green, below pale green to whitish, the two sep-
arated by a bicolored orange or brown (below) and white (above) (males) or white only or absent
(females) ventrolateral stripe, which occupies the whole of the outermost scale row and a portion
of the second row; ventrals 155-165 (—170); subcaudals (58—) 61—68, all paired; hemipenes short,
spinose beyond bifurcation. Total length 750 mm, tail length 145 mm.
DISTRIBUTION.— MYANMAR (Map p. 462):Chin State (Haka); Kachin State. ELSEWHERE:
China (Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi,
Jilin, Sichuan, Yunnan, Zhejiang); Taiwan; Vietnam (Lao Cai, Vinh Phu, and possibly Bac Thai, Da
Nang, Gia Lai, and Hoa Binh); Laos (after David et al. 2001:218).
HasitatT.— Recorded at 1206 m in the mountainous regions of Yunnan in Nujiang Langcang
Gorge alpine conifer and mixed forest. Das (2002) (as T: stejnegeri) reports this species at eleva-
tions up to 2845 m. Historical records for Myanmar place this species in subtropical forests (north-
ern triangle) and Chin Hills-Rakhine montane forest. Individuals have been recorded in bushes and
trees and on the ground in grass (Pope 1935), also from grassy slopes with shrubs as well as mixed
forest (Schleich and Kastle 2002).
REMARKS.— This species has been variously treated as a subspecies of T. stejnegeri and as a
distinct species allied to 7: stejnegeri. David et al. (2001) have referred all specimens previously
identified as 7: stejnegeri coming from Myanmar to 7: yunnanensis, and they have restricted 7: ste-
jnegeri to China, Taiwan, Vietnam and Laos. A specimen identified as 7: stejnegeri in the Natural
History Museum London, reportedly from Chin State, is considered by David et al. to be “an atyp-
ical specimen of 7: yunnanensis” (David et al. 2001:219). If David et al. are correct, then
Trimeresurus stejnegeri should probably be removed from the faunal list for Myanmar, although
the “stejnegeri’ clade is still represented in the country by T? yunnanensis and T. medoensis (q.v.).
Nonetheless, as we pointed out earlier (p. 409), there is a good possibility that 7. stejnegeri (sensu
stricto) will turn up in eastern Shan and Kachin States,* and for this reason, it is still listed here (see
above). On the other hand, we also concur with David et al. (2001:219) who argue, with justifica-
tion, that considerable work remains to be done and material examined from throughout the south-
ern ranges (from northeastern India to Vietnam) of the several recognized nominal species to
understand species boundaries within the “stejnegeri” clade.
LITERATURE CITED
Note: Not included in this section are full bibliographic citations for most pre 20th century literature
even though reference is made to them in the synonymies (e.g., Schneider, 1799, 1801; Cantor, 1839; Gray,
1834, 1842, 1849; Laurenti, 1768; Linnaeus, 1758, Schmidt, 1925, and others). Complete citations to these
will be found in the bibliography in GOLAy ET AL. 1993 (q.v.).
4 NB. Hallermann et al. (2002) identified one of Leonardo Fea’s specimens collected at “Mti. Carin [=Karen],
900-1000 m a.s.l. (ZMH R06267-8)” (Kayah State) as 7. stejnegeri. Kayah State is immediately south of Shan State.
450 | PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 24
ADLER, KRAIG, HOBART M. SMITH, SUSAN H. PRINCE, PATRICH DAVID, AND DAVID CHISZAR. 2000. Russell’s
viper: Daboia russelii, not Daboia russellii, due to classical Latin rules. Hamadryad, 25(2):83-85.
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British Museum, London. xiv + 727 pp., 25 pls.
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184-193 in P. Golay et al., Endoglyphs and Other Major Venomous Snakes of the World. A Checklist.
Azemiops S.A., Aire-Geneva, Switzerland.
CoGGER, HAROLD G. 1975 Sea Snakes of Australia and New Guinea. Pages 59-139 in W.A.Dunson, ed., The
Biology of Sea Snakes. University Park Press, Baltimore.
Cox, M. 1991. The Snakes of Thailand and their Husbandry. Kreiger Publishing, Malabar, Florida. 526 pp.
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Ltd., London, UK. 144 pp.
Davib, PATRICK, AND GERNOT VOGEL. 1996. The Snakes of Sumatra: An Annotated Checklist and Key with
Natural History Notes. Edition Chimaira, Frankfurt-am-Main, Germany. 260 pp., 33 col. photos.
Davib, PATRICK, AND HAIYAN TONG. 1997. Translations of recent descriptions of Chinese pitvipers of the
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pitviper, 7rimeresurus stejnegeri (Serpentes, Viperidae, Crotalinae), with the description of a new species
from Thailand. Russian Journal of Herpetology 8(3):205—222, col. figs. 1-7.
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HALLERMANN, JAKOB, NATALIA ANANJEVA, NIKOLAI ORLOV, AND FRANK TILLACK. 2002. Leonardo Fes’s histor-
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Mitteilungen der Hamburg Zoologisches Museum und Institut 99:139-153, figs. 1-6.
HOoGE, ALPHONSO R., AND S.A. ROMANO-HOGE. 1983. Notes on micro and ultrastructure of “Oberhautschen”
in Viperoidea. Mem. Instituto Butantan 44/45[1980/81]:81—-118.
INEICH, IVAN, AND PIERRE. LABOUTE. 2002. Sea Snakes of New Caledonia. Institute de Recherché pour le
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Verlagsbuchhandlung, Marburg.
Kraus, F., D.G. MINK, AND W.M. BRown. 1996. Crotaline intergeneric relationships based on mitochondrial
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key to the genus Trimeresurus. Copeia 1942(1):18—24, figs. 1-2.
McCartny, CoLin. 1993. Laticauda Laurenti, 1768. Pages 145-148 in P. Golay, et al., Endoglyphs and Other
Major Venomous Snakes of the World. A Checklist. Azemiops S.A., Aire-Geneva, Switzerland.
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HK OK OK OK OK OK 2K OK 2K OK 3K OK OK OK BK OK OK 2K OK 2K OK 2K OK 3K 2K
APPENDIX A
SNAKEBITE PROCEDURES IN MYANMAR
Prepared by Joseph B. Slowinski, Ph.D.
NOTICE: A word of caution. The following guidelines were formulated from recommenda-
tions made in the WHO/SEARO Guidelines for the Clinical Management of Snakebites in the
Southeast Asian Region (1999), written and edited by Dr. David Warrell, and from specific com-
ments made by Dr. David Warrell. However, recommendations and medical procedures undergo
constant review and revision; thus, the reader is cautioned to seek the latest information and guid-
ance from appropriate medical specialists before adopting any of the recommendations cited below.
In case of a bite from any species in the family Elapidae or Viperidae or a bad bite from a
Rhabdophis subminiatus, the bitten person must be transported to a hospital as soon as possible.
Do not allow traditional treatments; they will only delay transport to the hospital and can even
cause additional damage. Although a bite from a venomous snake should be considered a medical
emergency, the victim should be reassured and calmed as much as possible. During the transport
process to the hospital, the victim should move as little as possible—any muscle contractions will
increase systemic absorption of the venom. With treatment, the chances of dying from a venomous
snakebite are small.
The best hospital has the following:
1) antivenom:
2) epinephrine to treat anaphylaxis;
3) a system for assisted breathing in the case of neurotoxicity from an elapid bite;
4) treatment for acute kidney failure.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 453
Russell’s viper (Daboia russelii). Do not apply first aid, do not allow the patient to walk, pro-
ceed immediately to a hospital and request antivenom for Daboia russelii from the Myanmar
Pharmaceutical Industry. Severe systemic symptoms such as incoagulable blood, hemorrhage,
shock, and kidney failure can develop rapidly from bites by this species. At the hospital, blood
coagulability should be tested, which can be done by the 20-minute whole blood clotting test (see
below). The hospital should be prepared to treat the kidney failure that often results from bad bites
by this species.
Monocled cobra (Naja kaouthia). Do not apply first aid, do not allow the patient to walk, pro-
ceed immediately to a hospital and request antivenom for Naja kaouthia from the Myanmar
Pharmaceutical Industry. If breathing becomes difficult because of severe neurotoxicity—whether
or not antivenom has been administered—the patient should be placed on assisted breathing.
Neuromuscular transmission can often be dramatically improved with an anticholinesterase drug
such as neostigmine or edrophonium. Atropine should be given as well.
Spitting cobra (Naja mandalayensis). Do not apply first aid, do not allow the patient to walk,
proceed immediately to a hospital. Do not accept antivenom. If breathing becomes difficult because
of severe neurotoxicity (this species may or may not cause neurotoxicity; there is no published
information on the clinical effects of its venom), the patient should be placed on assisted breath-
ing. Neuromuscular transmission can often be dramatically improved with an anticholinesterase
drug such as neostigmine or edrophonium. Atropine should be given as well. There is anecdotal
evidence that the venom of this species can cause severe local effects, including necrosis.
In the case of spitting cobra venom spat in the eyes, the eyes should be immediately flushed
with generous amounts of water. Do not accept the traditional treatment of tamarind leaf juice in
the eyes. Apply topical antibiotic unless corneal abrasions can be excluded by fluorescein staining
or slit lamp examination.
King cobra (Ophiophagus hannah). Apply a compression bandage to the bitten limb, do not
allow the patient to walk, proceed immediately to a hospital. Do not accept antivenom, unless they
have antivenom specifically made for king cobras in India or Thailand. If breathing becomes dif-
ficult because of severe neurotoxicity—whether or not antivenom has been administered—the
patient should be placed on assisted breathing. Neuromuscular transmission can often be dramati-
cally improved with an anticholinesterase drug such as neostigmine or edrophonium. Atropine
should be given as well.
Elapid seasnake (Hydrophis, Lapemis, other genera). Apply a compression bandage to the
bitten limb, do not allow the patient to walk, proceed immediately to a hospital. Do not accept
antivenom. If breathing becomes difficult because of severe neurotoxicity, the patient should be
placed on assisted breathing. Neuromuscular transmission can often be dramatically improved with
an anticholinesterase drug such as neostigmine or edrophonium. Atropine should be given as well.
Note: local symptoms from bites by this group may be minimal and should not be taken as a sign
of no envenomation.
Krait (Bungarus). Apply a compression bandage to the bitten limb, do not allow the patient
to walk, proceed immediately to a hospital. Do not accept antivenom. If breathing becomes diffi-
cult because of severe neurotoxicity, the patient should be placed on assisted breathing.
Neuromuscular transmission can often be dramatically improved with an anticholinesterase drug
such as neostigmine or edrophonium. Atropine should be given as well. Note: local symptoms from
bites by this genus may be minimal and should not be taken as a sign of no envenomation.
Coral snake (elapid genera Maticora (= Calliophis), Calliophis, Sinomicrurus). Do not
454 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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apply first aid, do not allow the patient to walk, proceed immediately to a hospital. Do not accept
antivenom. If breathing becomes difficult because of severe neurotoxicity, the patient should be
placed on assisted breathing. Neuromuscular transmission can often be dramatically improved with
an anticholinesterase drug such as neostigmine or edrophonium. Atropine should be given as well.
Vipers (other than Daboia) (Ovophis, Protobothrops, Trimeresurus). Any viperid other than
the Russell’s viper (e.g., Trimeresurus). Do not administer first aid; proceed immediately to a hos-
pital, do not accept antivenom.
Red-necked keelback (Rhabdophis subminiatus). If the bite was bad (1.e., the snake chewed
the skin for more than several seconds), do not administer first aid, proceed immediately to a hos-
pital, do not accept antivenom. The blood should be tested for incoagulability. Note: the bad symp-
toms from this species may take more than a day to develop.
Antivenom Rules
1) Antivenom should only be administered if there are signs of local or systemic envenoma-
tion.
2) Before antivenom is injected, epinephrine must be available to counter anaphylaxis. If the
need arises to inject epinephrine, an antihistamine should also be applied to neutralize histamine
release during the allergic reaction.
3) Under no conditions should you allow antivenom from a species other than the one that bit
the victim to be administered.
4) Under ordinary circumstances, do not accept antivenom made in other countries, except
possibly in case of a bad king cobra bite.
5) The doctor should monitor the patient for adverse reactions for at least one hour after
antivenom is administered.
Definitions and Explanations
20-minute whole blood clotting test. A very simple test to perform. Place a few ml of blood
into a glass container; leave undisturbed for 20 minutes; tip the vessel once. If the blood is still liq-
uid and runs out, the patient has incoagulable blood, a sign of systemic poisoning.
Anaphylaxis. A potentially life-threatening allergic reaction to foreign molecules character-
ized by a dramatic drop in blood pressure. With snakebite, anaphylaxis can develop in two ways:
in response to the venom or in response to the antivenom. When antivenom is administered it is
important for the hospital to have epinephrine nearby to counteract a possible anaphylactic reac-
tion.
Neurotoxicity. The venoms of cobras, kraits, and sea snakes are dominated by neurotoxins,
compounds that adversely affect the nervous system. Symptoms of neurotoxic poisoning include
loss of muscle control, which often is manifested by drooping eyelids and loss of muscle tone in
other facial features. The major effect — and the deadly one — is the paralysis of the diaphragm
resulting in the inability to breathe. This is the reason for the necessity of assisted breathing mech-
anisms.
Antivenom. Antivenom is generally manufactured in horses. A large dose of antivenom caus-
es the body to react to the serum and so severely that the body’s immunological reaction is so
strong as to cause shock and possibly death from the antivenom. Epinephrine counteracts the
immunological reaction and reduces shock. Antivenom is dangerous for the reason described above
and is highly species specific. Taking an antivenom for another snake species has no medical value
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 455
because it will not counteract the venom of a different species, and it threatens the snakebite vic-
tim because of the potential immunological reaction to the serum. The antivenoms produced in
Myanmar use venom from Myanmar snakes. Because the venoms from the same species in differ-
ent geographic areas may differ, it is essential that you use Myanmar antivenoms to ensure that the
antivenom works to counter the venom injected by a Myanmar species. The Myanmar
Pharmaceutical Industry manufactures antivenom against bites from Russell’s vipers and monocled
cobras.
Compression bandage. The above recommendations for and against the use of compression
bandages are based on a consideration of a venom’s potential to cause local tissue damage versus
its potential to cause life-threatening systemic symptoms. A compression bandage should be
applied as tightly as for a sprained ankle. It should not be applied so tightly that it eliminates the
bitten limb’s pulse. A compression bandage applied to a bitten limb will retard the venom’s absorp-
tion by the body, but traps venom in that compartment running the risk of amplified local tissue
damage. The venom of kraits is known to be highly toxic, yet does not cause serious local tissue
damage. Hence, there is little risk of tissue damage by applying the bandage, only the benefit of
retarding venom spread until a hospital is reached. On the other hand, the venom of snakes of the
genus 7rimeresurus have the potential to cause severe local tissue damage, which will be made
much worse by the application of a compression bandage.
HK AE AK AK AE AK BK HK AK AE AK AK AK IC AK KK AK IKK OK IK AK OK
MAPS
Locations of dangerously venomous
snake materials collected by
Myanmar Herpetological Survey
teams 1999-2003
with additional localities taken
from literature records
Note: Maps are digitally rendered using ArcGIS® (ESRI) and
source material from U.S. Geological Survey (USGS), National
Imagery and Mapping Agency (NIMA), and Australian Centre of the
Asian Spatial Information and Analysis Network (ACASIAN).
Inspiration for the Myanmar Ecosystem map came from
reading Olson, David M., et al. 2001. Terrestrial ecoregions of the
World: A new map of life on Earth. BioScience 51(11).
All maps were prepared by Michelle S. Koo, Biogeographic
Information Systems Manager, Department of Herpetology,
California Academy of Sciences
456
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Venomous Snake
Collection Localities
by the
Myanmar
Herpetological Survey
1999 - 2003
ANDAMAN
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LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR
Myanmar
coastal
rain forest
BA Y Irrawaddy
ah fepseterind
_ South Thailand
d semi-evergreen
rain forest »; a
Myanmar Coast
mangrove
) ANDAMAN
ie
100
Z kilometers
© 2003 California Academy of Sciences
Ecoregions of Myanmar
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© 2003 California Academy of Sciences
Myanmar Herpetological Survey
Localities
ANDAMAN
Bungarus bungaroides* SEA
Bungarus fasciatus
Bungarus flaviceps*
Bungarus magnimaculatus
Bungarus wanghaotingi
* locality from literature; see checklist.
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 459
BAY
(ou
BENGAL
© 2003 California Academy of Sciences
Myanmar Herpetological Survey
Localities
ANDAMAN
Calliophis maculiceps SEA
Naja kaouthia
Naja mandalayensis
Ophiophagus hannah
Sinomicrurus macclellandi
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Myanmar Herpetological Survey
Localities
ANDAMAN
Enhydrina schistosa
Hydrophis ornatus
Lapemis hardwickii
Laticauda laticaudata
ek
Laticauda colubrina
osnjodiytY ¥
LEVITON ET AL.: DANGEROUSLY VENOMOUS SNAKES OF MYANMAR 461
BAY
OF
BENGAL
© 2003 California Academy of Sciences
Myanmar Herpetological Survey
Localities
ANDAMAN
Daboia russelii SEA
Ovophis monticola
Protobothrops jerdonii
Protobothrops kaulbacki
Protobothrops mucrosquamatus
462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Localities
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Trimeresurus albolabris SEA
Trimeresurus erythrurus
Trimeresurus medoensis
Trimeresurus popeiorum
Trimeresurus purpureomaculatus
Trimeresurus yunnanensis
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25, pp. 463-498, 20 figs., 4 tables November 14, 2003
Descriptions of Seven New Cyrtodactylus (Squamata:
Gekkonidae) with a Key to the Species of Myanmar (Burma)
Aaron M. Bauer!
Department of Biology, Villanova University
800 Lancaster Avenue, Villanova, Pennsylvania 19085
e-mail: aaron.bauer@ villanova.edu
Seven new species of the gekkonid lizard genus Cyrtodactylus are described on the
basis of material collected by the Myanmar Herpetological Survey. These include
four small to moderately sized species with relatively short digits and three larger
species with long, slender digits. Among the smaller forms two are allied to C.
khasiensis; one from the Ayeyarwady delta and adjacent lowlands, the other from
higher elevations in the Chin Hills. The remaining small species, from Alaungdaw
Kathapa National Park (Sagaing Division) and Rakhine Yoma Elephant Range
(Rakhine State) are probably most closely allied to C. consobrinoides. The large-bod-
ied forms include the apparent sister species of C. slowinskii, from far northwestern
Myanmar, and two other, distinctive species from Mon State and Shan State, respec-
tively. A dichotomous key to the sixteen species of Cyrtodactylus known from
Myanmar is presented. The discovery of seven new species of the genus suggests that
each hill range, as well as isolated peaks, may be expected to harbor endemic species
of geckos. The collections of the Myanmar Herpetological Survey in areas to the west
of the Ayeyarwady River complement historical collections derived chiefly from
areas to the south (the former Lower Burma) and to the west of the central arid zone.
Myanmar has long been recognized as a bridge region connecting the largely herpetofaunally
discrete areas of peninsular India and southeast Asia (Theobald 1868). Despite early collecting
activity in central and northern Burma (e.g., by Leonardo Fea, see Hallermann et al. 2002) as well
as in Tenasserim (Tanintharyi Division), along the Thai border the majority of the country has
remained poorly documented herpetologically (Inger 1999; Slowinski and Wiister 2000). Although
its reptile fauna has been recognized as being relatively rich, it is not noted for its endemism. The
recent series of expeditions conducted by the Myanmar Herpetofaunal Survey, a joint program of
the Myanmar Nature and Wildlife Conservation Division, Forest Department, the California
Academy of Sciences, and the Smithsonian’s National Museum of Natural History, with support
from the National Science Foundation, has, however, revealed a variety of new species, many
apparently endemic to Myanmar (e.g., Slowinski and Wiister 2000; Bauer 2002). Among the
lizards, one of the most interesting groups in this regard appears to be “bent-toed” geckos of the
genus Cyrtodactylus.
Despite the taxonomic revision and allocation of bent-toed geckos to a number of putatively
monophyletic and geographically cohesive groups by previous authors (Underwood 1954;
Szczerbak and Golubev 1977, 1984, 1986; Kluge 1983), Cyrtodactylus remains a large and
! Research Associate, Department of Herpetology, California Academy of Sciences
463
464 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
unwieldy group of approximately 70 species distributed from South Asia through the Indo-
Australian Archipelago as far as the Solomon Islands (Bauer and Henle 1994). Kluge (1991, 1993,
2001) recognized two genera of tropical Asian bent-toed geckos: Cyrtodactylus and Geckoella, the
latter occurring only in Peninsular India and Sri Lanka. This division has, however, not been uni-
formly accepted (e.g., ROsler 2000; Bauer 2002), and I here tentatively consider Geckoella as a
subgenus of Cyrtodactylus pending a phylogenetic analysis of the group as a whole or, alternative-
ly, evidence that Geckoella does not render remaining Cyrtodactylus paraphyletic.
Bauer (2002) reviewed the Cyrtodactylus of Myanmar and described two new taxa, bringing
the number of species recorded for the country to ten. A record of C. pulchellus from Myanmar
(Das and Lim 2000), however, has subsequently been revealed as unverified (I. Das, pers. com-
mun., 28 June 2003), leaving nine Cyrtodactylus confirmed for the country. This includes C.
rubidus, an Andaman Islands endemic, presumably present in the Cocos Group, the only part of the
Andamans under the administration of Myanmar (fide Hundley 1964), and the mainland species C.
brevidactylus, C. consobrinoides, C. feae, C. khasiensis, C. oldhami, C. peguensis, C. slowinskii,
and C. variegatus. Bauer (2002) also signaled the existence of two additional, undescribed species
based on unworked material from the Myanmar Herpetological Survey. Examination of this mate-
rial, as well as specimens collected subsequently, reveals the existence of seven new species of
Cyrtodactylus from Myanmar, for a total of 16 species. This is comparable to the 14 species now
known from the much more intensively surveyed neighboring country of Thailand (Bauer et al.
2003).
MATERIALS AND METHODS
The following measurements were taken with Brown and Sharpe Digit-cal Plus digital calipers
(to the nearest 0.1 mm): snout-vent length (SVL; from tip of snout to vent), trunk length (TrunkL;
distance from axilla to groin measured from posterior edge of forelimb insertion to anterior edge
of hindlimb insertion), crus length (CrusL; from base of heel to knee); tail length (TailL; from vent
to tip of tail), tail width (TailW; measured at widest point of tail); head length (HeadL; distance
between retroarticular process of jaw and snout-tip), head width (HeadW; maximum width of
head), head height (HeadH; maximum height of head, from occiput to underside of jaws), ear
length (EarL; longest dimension of ear); forearm length (ForeaL; from base of palm to elbow);
orbital diameter (OrbD; greatest diameter of orbit), nares to eye distance (NarEye; distance
between anteriormost point of eye and nostril), snout to eye distance (SnEye; distance between
anteriormost point of eye and tip of snout), eye to ear distance (EyeEar; distance from anterior edge
of ear opening to posterior corner of eye), internarial distance (Internar; distance between nares),
and interorbital distance (Interorb; shortest distance between left and right supraciliary scale rows).
Unless otherwise stated, counts and measurements made on right side of specimens.
Scale counts and external observations of morphology were made using a Nikon SMZ-10 dis-
secting microscope. Preserved specimen photographs were taken with a Nikon CoolPix 990 digi-
tal camera. Radiographic observations were made using a Faxitron closed cabinet x-ray system.
Comparisons were made with museum material in the collections of the California Academy of
Sciences (CAS), National Museum of Natural History (USNM), Museum of Comparative Zoology
(MCZ), Institut Royal des Sciences Naturelles de Belgique (IRSNB), and Chulalongkorn
University Museum of Zoology (CUMZ), as well as original published descriptions and descrip-
tions provided in broader faunal and taxonomic treatments (e.g., Annandale 1913; Smith 1935;
Szczerbak and Golubev 1986; Hikida 1990; Ulber 1993; Darevsky and Szczerbak 1997; Das 1997;
Das and Lim 2000; Bauer 2002; Bauer et al. 2002).
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 465
Geographic coordinates and elevation were recorded by the original collectors from a Garmin
12 GPS (datum WGS 84).
SYSTEMATICS
Reptilia: Squamata: Gekkonidae
Cyrtodactylus annandalei Bauer, sp. nov.
Figs. 1-2
HoLotyPe.— California Academy of Sciences (CAS) 215722 (Field number JBS 4995), adult
male; Gon Nyin Bin Camp, Alaungdaw Kathapa National Park, Monywa District, Sagaing
Division, Myanmar (22°14’51.1”N, 94°37'19.3”E); collected by Thin Thin, San Lwin Oo, Hla Tun,
Z.T. Aung, and T.T. Soe, 6 July 2000.
PARATYPES.— USNM 559840 (Field number JBS 5066), adult female; Pwint Kyi Camp,
Alaungdaw Kathapa National Park, Monywa District, Sagaing Division, Myanmar (22°14’39.8”N,
94°37’49.8”E); same collectors as holotype, 11 July 2000. CAS 215757 (Field number JBS 5077),
adult female; Pwint Kyi Camp, Alaungdaw Kathapa National Park, Monywa District, Sagaing
Division, Myanmar (22°14’39.8”N, 94°37’49.9”E); same collectors as holotype, 12 July 2000.
ETYMOLOGY.— The specific epithet is a patronym honoring Thomas Nelson Annandale
(1876-1924), founder and director of the Zoological Survey of India. Annandale described numer-
ous reptiles, including geckos from tropical Asia, especially India and Burma. In particular, he
(Annandale 1905b) described Gymnodactylus consobrinoides, the species most closely resembling
C. annandalei, from Tavoy (Tavai). The name is masculine and is formed in the genitive case.
DEFINITION.— A small sized Cyrtodacty- aS —
lus, snout-vent length 49-55 mm; body rela-
tively slender, limbs, and especially digits,
short; one pair of enlarged postmental scales in
broad contact behind mental; dorsum relatively
smooth textured, with 16-18 rows of small,
keeled tubercles; 43 ventral scales across mid-
body; no precloacal groove, 11-12 precloacal
pores in a single series in both males and
females, 10—11 femoral pores in males, separat-
ed from precloacal series by a diastema. Ten
subdigital lamellae beneath 4' toe of pes distal
to digital inflection. Subcaudal scalation of
original tail with alternating rows forming wide
transverse plates. Dorsal pattern of 6 dark bands
(one each on occiput and nape, four on trunk
and sacrum) or rows of transversely arranged
spots, with narrow whitish borders. Dorsum of
head patternless. Tail with alternating light and
dark banding.
DESCRIPTION (based on holotype, CAS
215722).— Adult male with midventral inci-
sion from tissue removal. Snout-vent length Figure 1. Holotype of Cyrtodactylus annandalei, sp.
49.0 mm. Head relatively long (HeadL/SVL nov. (CAS 215722) from Alaungdaw Kathapa National Park,
Myanmar. Note the relatively short digits, patternless head
ratio 0.28), moderately wide (HeadW/HeadL — dorsum, and distinct dorsal pattern. Scale bar = 10 mm.
466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
ratio 0.61), not depressed (HeadH/HL ratio 0.42), distinct from neck. Lores and interorbital region
inflated, canthus rostralis not particularly prominent. Snout moderately short (SnEye/HeadL ratio
0.39); longer than eye diameter (OrbD/SnEye ratio 0.60); scales on snout and forehead rounded,
granular, flattened to slightly conical, becoming heterogeneous posteriorly; scales on snout larger
than those on occipital region. Eye small (OrbD/HeadL ratio 0.23); pupil vertical with crenelated
margins; supraciliaries short, without projecting spines. Ear opening oval, moderately large
(EarL/HeadL ratio 0.07); eye to ear distance greater than diameter of eyes (EyeEar/OrbD ratio
1.32). Rostral approximately half as deep (1.2 mm) as wide (2.4 mm), incompletely divided dor-
sally by weakly developed rostral groove; two enlarged supranasals separated by two much small-
er internasals; rostral in contact with supralabial I, supranasals, and internasal; nostrils circular,
each surrounded by supranasal, rostral, first supralabial, and three postnasals, rostral contact of nar-
ial border extensive; narial flap partially occludes posterior third of nostril; two to three rows of
scales separate orbit from supralabials. Mental subtriangular, much wider (2.2 mm) than deep (1.6
mm); one pair of enlarged postmentals, each 60% size of mental, in broad contact with one anoth-
er medially, bordered anterolaterally by first infralabial, posterolaterally by enlarged lateral chin-
shield, and posteriorly by 2—3 chin granules. Infralabials bordered medially by 1-3 rows of
enlarged scales. Supralabials (to midorbital position) 7 (left) —8 (right); enlarged supralabials to
angle of jaws 10; infralabials 9; interorbital scale rows across narrowest point of frontal 12.
Body relatively slender, relatively short (TrunkL/SVL ratio 0.40) with ventrolateral folds
indistinct. Dorsal scales small, granular, conical, with regularly arranged small tubercles extending
from posterior border of orbits and temporal region on to tail base; each tubercle rounded, bearing
a single prominent keel; tubercles becoming smaller and less prominently keeled on flanks; largest
keeled tubercles in approximately 16 regular rows at midbody. Ventral scales much larger than dor-
sal, smooth, subimbricate laterally, strongly imbricate across midventer; somewhat larger than else-
where along ventral midline of chest and abdomen, and especially in precloacal region; midbody
scale rows across belly to edge of flanks (as demarcated by dorsal color pattern) 43; gular region
with homogeneous scalation. Precloacal pores in a single continuous series of 11; bordered poste-
riorly by a row of greatly enlarged poreless scales. Femoral pores present in continuous rows of 10
(left) — 11 (right), separated from precloacal pores by a diastema of 5 scales; no precloacal groove.
Scales on palm and sole smooth, rounded; scales on dorsal aspects of limbs smooth, subimbricat-
ing, heterogeneous but without tubercles.
Fore- and hindlimbs relatively short, stout; forearm short (ForeaL/SVL ratio 0.14); tibia mod-
erate (CrusL/SVL ratio 0.17); digits short, strongly inflected at basal interphalangeal joints, all
bearing robust, slightly recurved claws; subdigital lamellae rounded, smooth, without scansorial
surfaces; lamellae distal to much enlarged scale at basal digital inflection and not including ventral
claw sheath: 6—7—8—8-8 (manus), 8-8—10—10—10 (pes), proximal 1—2 fragmented on most digits;
enlarged basal lamellae (> twice size of palmar scales) to and including enlarged scale at basal
inflection: 3—4—5—5—4 (manus), 3—5—6—7-—S (pes); interdigital webbing absent. Relative length of
digits (manus; measurements in mm in parentheses): III (3.4) ~ IV (3.4) > V (2.7) ~ II (2.6) >I
(2.4); (pes): IV (4.5) > V (4.2) > III (3.8) > II (G.3) > I (2.3).
Tail original, slightly longer than body (TailL/SVL ratio 1.04), slender, tapering; divided into
indistinct segments, each approximately 7 dorsal scale rows in length; two rows of enlarged, keeled
tubercles positioned paravertebrally on tail base only, remaining dorsal caudal scales small,
smooth, rectangular; subcaudal scales larger, smooth, imbricate; subequal basally, distally with
alternating midventral scales enlarged into transverse plates. Two enlarged, smooth, conical post-
cloacal spurs on each side of tailbase.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3—4—5-3 for
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 467
New Species
Cyrtodactylus
@ C aequalis
[]_ C. annandalei
BBC ayeyarwadyensis
Z\_ C. chrysopylos
A C. gansi
@ C.uwsselli
© C. wakeorum
all
C. chrysopylo
C. ayeryarwadyensis
type loc.
C. wakeorum
type loc.
FIGURE 3. Map of Myanmar illustrating the distribution of seven new species of Cyrtodactylus: C. aequalis (red circle),
C. annandalei (yellow square), C. ayeyarwadyensis (red square), C. chrysopylos (yellow triangle), C. gansi (green trian-
gle). C. russelli (blue circle), and C. wakeorum (yellow circle). Map prepared by Michelle S. Koo, California Academy of
Sciences.
468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
manus and 2-34—5-4 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 32 post pygal caudal vertebrae in original tail in holo-
type (32 in larger paratype; 5.5 in smaller paratype with regenerate tail). One pair of slender, cres-
centic cloacal bones in male holotype (absent in female paratypes). Endolymphatic sacs not
enlarged extracranially. Subadult female with significant areas of incomplete ossification at joints
of long bones.
Coloration (in preservative). Base color a pale brown. Boldly marked with chocolate brown
bands, each outlined by a thin cream to whitish border 1—2 scale widths in thickness. Occipital band
extending anteriorly to orbit and under eye to loreal region; continues indistinctly to rostral.
Dorsum of head patternless. Nuchal collar extending anteriorly to posterior border of ear. Four
additional bands between shoulders and sacrum. Ventrolateral pale brown spaces between body
bands with large, chocolate brown spots. Pattern bilaterally symmetrical on head and nape, some-
what asymmetrical more posteriroly. Forelimbs mottled, with an indistinct dark line along preaxi-
al surface of humerus; hindlimbs boldly marked with spots and oblong markings like those on
flanks. Hindlimb insertions with large chocolate brown blotches at posterior border of thighs.
Venter cream tinged by the light brown speckling of individual scales. Tail mottled gray-brown
with darker irregular transverse markings, except on enlarged imbricate median scales. Tail with
alternating chocolate brown and light brown to cream banding. Paler bands wider than darker, 12
dark bands from cloaca to tail tip. Tail
venter with scattered dark pigmentation,
especially distally. Dorsal color pattern
extends onto ventrum only at tail tip.
In life, dorsum distinctly darker than
flanks. Pale borders of dark spots yellow-
ish-cream (Fig. 2).
VARIATION.— Comparative mensural
data for the holotype and paratypes are
presented in Table 1. Both paratypes simi-
lar to holotype in most respects except as
noted. CAS 215757: adult female with
midventral incision from tissue removal FIGURE 2. Living specimen (paratype) of Cyrtodactylus annan-
dalei (CAS 215757). Photo by Hla Tun.
18 rows of keeled tubercles across mid-
body. 12 tiny precloacal pores and 4 (left) and 10 (right) weakly developed femoral pores. Nuchal
collar fragmented anteriorly; dark markings present above forelimb insertions, diffuse dark line
absent from preaxial border of humerus; 11 dark bands on original tail. CAS 215749: subadult
female with midventral incision from tissue removal. 18 rows of keeled tubercles across midbody.
Broken row of 8 tiny precloacal pores and 7 weakly developed femoral pores on each side. Dorsal
pattern highly fragmented into spots rather than bands; 4 spots in a transverse row reprepresenting
each posterior body band; hindlimbs with only a few small spots; infralabials brownish; regenerat-
ed tail mottled, without banding.
DIAGNOSIS.— Cyrtodactylus annandalei may be distinguished from all congeners on the basis
of its relatively short digits, series of 8-12 precloacal pores separated by a diastema from 4-12
femoral pores on each thigh, no precloacal groove or enlarged femoral scales, dorsal tubercles
small, in 16—18 longitudinal rows, ventral scales in 43 rows, alternate subcaudal scales enlarged
transversely, and dorsal color pattern, including occipital, nape, and four additional dark body
bands (or series of spots), and patternless head dorsum.
The new species is superficially most similar to C. consobrinoides (Annandale, 1905a;
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 469
TABLE |. Mensural data for the type series of Cyrtodactylus annandalei, sp. nov.
Abbreviations as in Materials and Methods section; all measurements in mm.
eAsais722 CAS 215757 CAS 215749
Holotype paratype paratype
ar idea Sales ae aaa female —sfemmaile:~
SVL 49 5503 Shell
ForeaL Uf 8.2 6.7
CrusL 8.4 OS) 8.6
TailL (entire) 50.8 55.5 36.4
TailL (portion regenerated) — — 22.8
TailW 4 é 4.3
TrunkL 19.7 22.6 22.9
HeadL 135) 2 13.8
HeadW 8.3 : 8.1
HeadH 5.6 6 4.7
OrbD 32 3S Se)
EyeEar 4.2 4.6 3.9
SnEye a3 5.6 ee
NarEye 3.6 3) 355
Interorb D3) D2, 4.5
EarL 1 1S) eg)
Internar Dep) 2 1.9
Annandale 1905b), from which it differs in having a smaller number of thicker dark crossbands
(6-7 vs. 8-9), no enlarged femoral scales, femoral pores present and separated from precloacal
scales by a diastema, a larger number of ventral scale rows (43 vs. 20-30, although this count is
somewhat arbitrary in the absence of ventrolateral folds as lateral landmarks for such counts), and
a patternless head dorsum. It also lacks the light borders around the dorsal markings illustrated by
Annandale (1913) in his redescription of the holotype of C. consobrinoides. A specimen from
Molmein (Mawlamyine), provisionally referred to C. consobrinoides by Smith (1935) has 40 ven-
tral scale rows and only six dorsal bands and may be referable to C. annandalei, although the huge
distance (ca. 680 km) between Mawlamyine and the type locality make this unlikely. Comparisons
with other species are provided following the description of all new taxa.
DISTRIBUTION.— Cyrtodactylus annandalei is known only from Alaungdaw Kathapa National
Park in the Sagaing Division of north central Myanmar (Fig. 3). The most similar species to this
form, C. consobrinoides is known only from southern Myanmar, with records from Tavoy (Dawei,
Tanintharyi State) (Annandale 1905a, 1905b, 1913) and Molmein (Mawlamyine, Mon State)
(Smith 1935; but see above). The species is sympatric with the much larger and recently described
C. slowinskii as well as an unidentified species related to C. khasiensis (USNM 548140). The type
locality is in the central dry zone of Myanmar, between the Rakhine Yoma (Arakan Yoma) of the
Indo-Burman Range in the west and the extensive montane areas east of the Sittaung River.
Cyrtodactylus ayeyarwadyensis Bauer, sp. nov.
Figs. 4-6
Ho.LotyPe.— CAS 226154 (Field number JBS 8689), adult male; Khoko Gwe Creek,
470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
Rakhine Yoma Elephant Range, Gwa Township, Rakhine State, Myanmar (17°43’48.3°N,
94°39'02.7’E); collected by J.B. Slowinski, GO.U. Wogan, Kyi Soe Lwin and Hla Tun, 28 April
2001.
PARATYPES (8 specimens).— CAS 226153 (Field number JBS 8688), adult male; same data as
for holotype. CAS 226147 (Field number JBS 8346), adult female; Kanthaya, Gwa Township,
Rakhine State, Myanmar (17°43’24.9’N, 94°32’08.3”E); collected by J.B. Slowinski, GO.U.
Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein and Hla Tun, 22 April 2001. CAS
226149 (Field number JBS 8519), adult male; Kyat Stream, Rakhine Yoma Elephant Range, Gwa
Township, Rakhine State, Myanmar (17°42’14.0’N, 94°38’54.3”E); collected by J.B. Slowinski,
G.O.U. Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein and Hla Tun, 26 April
2001. USNM 559837 (Field number JBS 8635), adult male; locality and collectors as for CAS
226149, 27 April 2001. CAS 226152 (Field number JBS 8638), adult female; data as for CAS
226150; USNM 559838 (Field number JBS 8637), adult female; data as for CAS 226150. CAS
226156 (Field number JBS 8782), adult female; Kanthaya, Gwa Township, Rakhine State,
Myanmar (17°43'14.6’N, 94°32’04.9”E); collected by J.B. Slowinski, GO.U. Wogan, Htun Win,
Thin Thin, Kyi Soe Lwin, Awan Khwi Shein and Hla Tun, 2 May 2001. CAS 216506 (Field num-
ber JBS 7467), adult male; Elephant Camp, Rakhine Yoma Elephant Range, Gwa Township, Than
Dawe District, Rakhine State, Myanmar (17°39’01.1’N, 94°38°39.2”E); collected by J.B.
Slowinski and Htun Win, 30 November 2000.
ADDITIONAL MATERIAL (16 specimens).— All specimens from Myanmar. CAS 226146 (Field
number JBS 8342); Kanthaya, Gwa Township, Rakhine State (17°43’24.9’N, 94°32’08.3”E); col-
lected by J.B. Slowinski, GO.U. Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein
and Hla Tun, 22 April 2001. CAS 226151 (Field number JBS 8636); Kyat Stream, Rakhine Yoma
Elephant Range, Gwa Township, Rakhine State (17°42’14.0’N, 94°38’54.3”E); collected by J.B.
Slowinski, G.O.U. Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein and Hla Tun,
27 April 2001. CAS 221934 (Field number JBS 8946); Rakhine Yoma Elephant Range, Gwa, Gwa
Township, Rakhine State (17°35’02.4”N, 94°40’44.2”E); collected by Htun Win, Thin Thin, Kyi
Soe Lwin and Awan Khwi Shein, 2 June 2001. CAS 221985 (Field number JBS 9296); Kyauk Win
Gyi Camp, Gwa Township, Rakhine State (17°53’59.9’N, 94°53’36.8”E); collected by Kyi Soe
Lwin, Awan Khwi Shein and Hla Tun, 9 June 2001. CAS 226155 (Field number JBS 8781), CAS
226157 (Field number JBS 8783), CAS 226158—226159 (Field numbers JBS 8792, 8794);
Kanthaya, Gwa Township, Rakhine State (17°43’14.6”N, 94°32’04.9’E); collected by J.B.
Slowinski, GO.U. Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein and Hla Tun, 2
May 2001. CAS 216526 (Field number JBS 7496); Elephant Camp, Rakhine Yoma Elephant
Range, Gwa Township, Than Dawe District, Rakhine State (17°38’58.3”N, 94°38’14.8”E); collect-
ed by J.B. Slowinski and Htun Win, 1 December 2000. CAS 216446 (Field number JBS 7363);
vicinity of Kanthaya Beach, Gwa Township, Rakhine State (no coordinates recorded); collected by
J.B. Slowinski and Htun Win, 27 November 2000. CAS 212459 (Field number JBS 4647); vicini-
ty of Mwe Hauk Village, Myaung Mya Township, Ayeyarwady Division (16°16'29.4’N,
94°46'04.0’E); collected by J.B. Slowinski, GR. Zug, R.S. Lucas and J.V. Vindum, 22 April 2000.
CAS 222812 (Field number JBS 11012); Mwe Hauk Village, Myaung Mya Township, Ayeyarwady
Division (16°16'34.8’N, 94°45’46.8”E); collected by GO.U. Wogan, Htun Win, Awan Khwi Shein,
Kyi Soe Lwin and Hla Tun, 20 January 2002.
In addition, the following specimens, although differing in minor aspects of coloration from
the types of C. ayeyarwadyensis, appear to be referable to this species. Further collecting in inter-
vening areas, however, may necessitate the reevaluation of these northern populations: CAS
223044 (Field number JBS 11324); Sa Byin Village, Taung Gok Township, Rakhine State
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 47]
(19°11’56.1’N, 94°11'56.1”E, 61 ft.); collected by GO.U. Wogan, Thin Thin, Kyi Soe Lwin, Awan
Khwi Shein and Hla Tun, 27 January 2002. CAS 223285 (Field number JBS 11691); Tha Byut
Stream, Ma Ei Ywa Ma Village, Taung Gok Township, Rakhine State (19°18’51.0’N,
94°09’06.5”E); collected by GO.U. Wogan, R.S. Lucas, Htun Win, Awan Khwi Shein and Kyi Soe
Lwin, 2 February 2002. CAS 223289 (Field number JBS 11698); Tha Byut Stream, Ma Ei Ywa Ma
Village, Taung Gok Township, Rakhine State (19°18’49.2’N, 94°09’08.8”E); collected by G.O.U.
Wogan, R.S. Lucas, Htun Win, Awan Khwi Shein and Kyi Soe Lwin, 2 February 2002. CAS
223339 (Field number JBS 11724); Tha Byut Stream, Ma Ei Ywa Ma Village, Taung Gok
Township, Rakhine State (19°18’51.7”N, 94°09’07.8”E); collected by G.O.U. Wogan, R.S. Lucas,
Htun Win, Awan Khwi Shein and Kyi Soe Lwin, 2 February 2002.
ETYMOLOGY.— The specific epithet is derived from the Ayeyarwady (Irrawaddy) River which
runs through Myanmar. The range of the species includes parts of the Ayeyarwady delta.
DEFINITION.— A moderate sized Cyrtodactylus, snout-vent length to 78 mm; body relatively
slender, limbs moderately long and digits relatively short; one pair of enlarged postmental scales,
in broad contact with one another behind mental; 22—24 rows of keeled, oblong dorsal tubercles;
32-37 ventral scales between ventrolateral folds; no precloacal groove, 10—28 precloacal pores in
a single series (or with scattered gaps of one poreless scale) in males only. Six widened subdigital
lamellae beneath basal phalanx of 4" toe of pes, 10 narrow lamellae beneath more distal phalanges
of same toe (in holotype). Subcaudal scalation without
enlarged midventral plates. Dorsal pattern of 9—11 (usually
10) transverse rows of rectangular brown blotches from
occiput to sacrum. Posterior border of each row usually
marked by white punctations or a narrow white band; tail
with alternating brown and white bands.
DESCRIPTION (based on holotype, CAS 226154).—
Adult male, SVL 63.4 mm. Head moderately long
(HeadL/SVL ratio 0.28), relatively wide (HeadW/HeadL
ratio 0.61), somewhat depressed (HeadH/HeadL ratio 0.35),
distinct from neck. Lores and interorbital region weakly
inflated, canthus rostralis not well developed. Snout moder-
ately long (SnEye/HeadL ratio 0.41); much longer than eye
diameter (OrbD/SnEye ratio 0.55); scales on snout and fore-
head rounded, granular, intermixed with scattered small
tubercles posteriorly; scales on snout much larger than those
on occipital region. Eye small (OrbD/HeadL ratio 0.22);
pupil vertical with crenelated margins; supraciliaries short,
blunt. Ear opening oval, obliquely oriented, large
(EarL/HeadL ratio 0.09); eye to ear distance greater than
diameter of eyes (EyeEar/OrbD ratio 1.21). Rostral approx-
imately 55% as deep (1.5 mm) as wide (2.8 mm), divided
dorsally by rostral groove; two enlarged supranasals separat- i
ed by a single, somewhat smaller, roughly hexagonal inter- Figure 4. Holotype of Cyrtodactylus
nasal; rostral in contact with supralabial I, supranasals, and @evarwadyensis, sp. nov. (CAS 226154)
internasal; nostrils oval, laterally oriented, each in broad ea RENAtD ome Ble paaL Range
3 : ownship, Rakhine State, Myanmar. Note
contact with rostral and also surrounded by supranasal, first the relatively short digits, paired rectangular
supralabial, and three postnasals; pigmented narial flap par- dorsal markings and white punctuations.
tially occludes posterior 2/3 of nostril; 3-4 rows of scales Scale bar = 10 mm.
472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
separate orbit from supralabials. Mental triangular, wider (2.8 mm) than deep (2.0 mm); one pair
of greatly enlarged postmentals, each approximately 30-35% size of mental; left and right post-
mentals in broad medial contact with no intervening granules, each member of pair bordered later-
ally by first infralabial and an enlarged lateral chinshield, the pair bordered posteriorly by 5 some-
what enlarged chin scales. Infralabials bordered by 1—3 rows of enlarged scales, largest anteriorly
and laterally. Throat scales small, rounded, granular. Supralabials (to midorbital position) 8 (left) —
10 (right); enlarged supralabials to angle of jaws 12 (left)—11 (right); infralabials 10; interorbital
scale rows across narrowest point of frontal bone 17.
Body moderately slender, relatively short (TrunkL/SVL ratio 0.40) with very weakly denticu-
late ventrolateral folds. Dorsal scales heterogeneous, mostly rounded to weakly conical granules
with pitted or rugose surfaces, intermixed with regularly arranged small (4-6 times granule size),
keeled, oblong tubercles extending from occipital region on to back and base of tail; tubercles on
nape more strongly conical and without keels; tubercles in approximately 22 longitudinal rows at
midbody; 50 tubercles in paravertebral row from occiput to mid sacrum . Ventral scales much larg-
er than dorsal, cycloid, imbricate to subimbricate; not enlarged under thighs or between precloacal
pores and vent; midbody scale rows across belly between ventrolateral folds 34; scales on throat
minute, granular, grading into larger scales on chest. Precloacal pores in a single series of 21, with
a gap of one poreless scale separating the distalmost two pores of the right side; no femoral pores;
no precloacal groove. Scales on palm and sole smooth, flattened; scales on dorsal aspects of
hindlimbs granular, conical, similar to dorsal scales, with larger, conical tubercles interspersed.
Dorsal scales of proximal forelimbs imbricate, without tubercles; scales of forearms heterogeneous
with scattered conical tubercles.
Fore and hindlimbs relatively slender; forearm (ForeaL/SVL ratio 0.16) and tibia (CrusL/SVL
ratio 0.18) relatively long; digits relatively short, strongly inflected at each joint, all bearing robust,
recurved claws; subdigital lamellae widened beneath basal phalanx; lamellae from first proximal
scansor greater than twice largest palm scale to basalmost digital inflection: 4~5—5—5—4 (manus)
and 3—5—5—6-—5 (pes); lamellae from basalmost digital inflection to toe tip, not including ventral
claw sheath: 7—8—8—7—7 (manus) and 7—10—8—10—10 (pes); one to several rows of small, non-
lamellar granules between basal and distal lamellar series; interdigital webbing present but weak-
ly developed. Relative length of digits (manus; measurements in mm in parentheses): IV (5.5) > III
64) > G7) > V 4:6) >1 G5): pes)? V'(7:0) SIV 6:6) > 164) SS OSlr as):
Original tail longer than body (TailL/SVL ratio 1.11), slender, cylindrical in cross section.
Scales arranged in regular segments; ventral scales rectangular, dorsal scales rounded at free mar-
gins. Eight circumferential rows per segment; basal segment with one row of 6 enlarged keeled
tubercles, each tubercle separated from next by 1-4 smaller scales; more distally tubercles are
unkeeled; tubercles absent on distal 3/4 of tail; posteriormost caudal scales narrow and elongate.
Subcaudal scales larger, 6 rows per segment; segments not strongly demarcated ventrally or poste-
riorly; no enlarged median plates. Cloacal spurs with 3 enlarged, smooth, rounded scales on each
side of tail base.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3—4-5-3 for
manus and 2—3—4—5-4 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 32 post pygal caudal vertebrae in original tail in holo-
type (32-34 in paratypes with complete tails, 0.5—21.5 in those with regenerated tails). One pair of
slender, crescentic cloacal bones in males at level of first to second pygal vertebrae (absent in
females). Endolymphatic sacs not enlarged extracranially.
Coloration (in preservative). Base color a medium brown marked by 10 transverse rows of
roughly rectangular dark brown patches from occiput to sacrum, each row consisting of a pair of
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 473
bold paravertebral markings and a pair of less well defined lateral markings. Each marking bolder
and more well-defined posteriorly and more diffuse anteriorly. Occipital markings fused to form a
complete band. Posterior border of each transverse row of rectangular patches marked by a series
of tiny white punctuations, each one to several scales in extent. Top of head medium brown with a
diffuse set of dark brown marks forming a semicircle between posterodorsal corners of orbits.
Canthal region crossed by a diffuse dark bar. A brown streak, bordered by cream, running from
behind orbit onto neck, bordered above and below by white punctuations. White punctuations also
scattered from insertion of forelimb to corner of mouth and along lower flanks. Limbs more-or-less
strongly barred, with alternating light and dark markings extending on to digits, lighter panes we
scattered white punctuations. Paravertebral marks fus-
ing on tail base to form complete dark bars, 12 such
bands along length of tail, including tail tip. First four
bands bordered posteriorly by white punctations and
alternating with narrower light bands. More posterior-
ly wide dark bands alternate with continuous narrow
white bands. Tail patterning extends on to venter, but
bands weakly demarcated anteriorly. Venter beige with FiGuRE 5. Living specimen of Cyrtodactylus
scattered dark pigment on all surfaces, darkest at later- ayeyarwadyensis, sp. nov. (CAS 226158) illustrat-
al margins of flanks, on neck, limbs, and around cloa- _ ins life coloration. Photo by Hla Tun.
ca. Palms and soles of feet dark.
Color in life similar to that in preservative
but with medium brown of body ranging from
straw to pinkish brown in some specimens.
Venter white with a pale pinkish suffusion on
flanks. Iris olive to greenish-gold. (Figs. 5—6)
VARIATION.— Comparative mensural data
for the holotype and paratypes are presented in
Table 2. Paratypes similar to holotype in most
respects except as noted. Scale rows between
ventrolateral folds at midbody 32—37. Rows of
dorsal tubercles at midbody 22-24. Precloacal @ RS Re tiie os a ale [ .
pores 10-28, generally continuous, but with a FIGURE 6. Head and forebody of living paratype of
single poreless scale dividing the series into 24 Cyrtodactylus ayeyarwadyensis, sp.nov. (CAS 216506)
fein aud™ (iehiypored scales CAS 216506, 4 cractt masa Hna) ie
pores absent in females. First supralabial scales
excluded from nostril in some specimens (CAS 216506, CAS 226156), nostril contacted by one
postnasal and a large crescentic nasal (CAS 216506, 226147, 226149, 226152), two postnasals and
a crescentic nasal (CAS 226156, USNM JBS 8637), or by an asymmetrical number of postnasals
(3 left, 4 right in CAS 226150, 4 left, 5 right in 226153). The largest of the paratypes, CAS 226152,
is a gravid female with two eggs clearly visible through the abdominal wall. Among the addition-
al specimens referred to C. ayeyarwadyensis, sizes range from 34.4 mm (CAS 212459, juvenile)
to 78.3 mm (CAS 221985, adult female).
Color pattern highly variable. Among the paratype series the number of pairs of dorsal mark-
ings was 10, as in the holotype, for most specimens, but ranged from 9 (CAS 216506, 226150,
226156) to 11 (CAS 226147). These markings may be evenly paired or partly phase shifted
between the left and right sides of the animal. The dark dorsal markings are especially strongly con-
trasting with respect to background color in CAS 216506 and 226149. The white punctuations may
474 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
TABLE 2. Mensural and meristic data for the type series of Cyrtodactylus ayeyarwadyensis, sp. nov.
Abbreviations as in Materials and Methods section; all measurements in mm.
226154 216506 226147 JBS8637 226156 226149 226150 226153 226152
| CAS CAS CAS USNM CAS CAS CAS CAS CAS
Rea Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
Sex male male female female female male male male female
SVL 63.4 67.6 64.1 65.9 65.5 66.6 62.1 65.4 71.8
ForeaL 9.8 10 9.4 9.8 9.5 10.1 9.9 9.6 11.8
CrusL 11.6 12 11.5 Ley 12.1 12.3 IES 10.7 13.1
TailL (total) 70.1 53.2 64.6 133 70.7 West 71 71.5 6.1
TailL (regen) — 43.3 13 — = = — M35) (broken)
TailW 6.9 6.3 58) 5.4 5.4 5.6 Dail 5.8 6.8
TrunkL 25.5 28.5 28.8 29 28 Die 28.3 26.7 34.2
HeadL 18 18.5 18.8 18.1 19 18.9 Nell 18.6 20.7
HeadW 11 1255 11.9 11.6 11.3 11.1 it 11.4 12.8
HeadH 6.3 6.8 6.4 6.9 7.3 6.9 6.3 6.9 8.5
OrbD 4 48 4.1 4.6 4.9 4.6 4.2 4.4 5
EyeEar 4.9 5.4 4.8 5.4 53 5 4.7 4.6 5.5
SnEye Je3 7.6 7 6.9 7.4 es) 7 7 8.2
NarEye Sal 4.8 49 5 4.9 all 4.5 49 5.8
Interorb 6.2 6.9 5.8 6.5 6.4 6.2 5 7.1 7.6
Eark 1.6 Dal 2A 1.6 1.6 1.9 1.8 28 Dell
Internar 2 Dal 2.4 7) ?) 1.8 1.8 1.9 Dal
Tubercle Rows 22 24 22 22 22 22 22 24 22
Ventral Scale Rows 34 34 34 36 34 37 34 32 36
Precloacal Pores 21 28 — = = 13 10 10 —
be greatly reduced or lacking almost all together (CAS 216506). Among the additional material
referred to this taxon pattern was even more variable, with several specimens from Rakhine State
(CAS 223285, 223289, 223339) with the white markings coalescent, forming distinct transverse
bands.
DIAGNOSIS.— Cyrtodactylus ayeyarwadyensis may be distinguished from all congeners on the
basis of its possession of short digits, a single series of precloacal pores (10—28) in males only,
absence of femoral pores and precloacal groove, 22—24 longitudinal rows of dorsal tubercles;
32-37 ventral scales between ventrolateral folds, subcaudal scalation without enlarged midventral
plates, and dorsal pattern of 9-11 (usually 10) transverse rows of rectangular brown blotches from
occiput to sacrum, usually bordered posteriorly by white punctations or a narrow white band.
Among other species from Myanmar it is most similar to C. khasiensis and a new species from
Chin State, described below. All are similar in size and bear a series of dark dorsal markings.
Cyrtodactylus ayeyarwadyensis differs from C. khasiensis, however, in having (in some speci-
mens) a greater number of precloacal pores (maximum 28 vs 14), rectangular dorsal blotches (vs a
more variegated pattern of alternating irregular light and dark bands, or even stripes; Hora 1926),
white dorsal punctuations or lines bordering dorsal blotches (in most cases), and narrow white tail
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 475
bands (vs subequal light and dark bands in C. khasiensis). It differs from the other new species in
both color pattern (rectangular dark markings and white punctuations vs narrow transverse bands)
and precloacal pore configuration (strongly angled and recessed into a shallow groove in the Chin
State species). Comparisons with other species are provided after the new species descriptions.
DISTRIBUTION.— Typical specimens of Cyrtodactylus ayeyarwadyensis are known from low
elevation in extreme southwestern Myanmar, west of the main channel of the Ayeyarwady
(Irrawaddy) River, specifically from Gwa Township in Rakhine State and Myaung Mya Township
in the Ayeyarwady Division. Additional specimens tentatively referred to C. ayeyarwadyensis
derive from Taung Gok Township, Rakhine State. This locality is considerably north of the
Ayeyarwady delta, but is also low-lying, to the west of the southern chain of the Rakhine Yoma
(Arakan Yoma) (Fig. 3).
Cyrtodactylus gansi Bauer, sp. nov.
Figs. 7-9
Ho.otype.— CAS 222414 (Field number JBS 8300); Che Stream, Min Dat Township, Min
Dat District, Chin State (21°2115.5”N, 93°56’13.3”E, 780 m.); collected by Htun Win, Thin Thin,
Kyi Soe Lwin, Awan Khwi Shein and Hla Tun, 3 April 2001.
PARATYPES.— CAS 222411 (Field number JBS 8234); Che Stream, Min Dat Township, Min
Dat District, Chin State (21°21’14.9”N, 93°56’08.3”E, 750 m.); collected by Awan Khwi Shein, 29
March 2001. USNM 559839 (Field number JBS 8258), CAS 226145 (Field number JBS 8260);
same locality as CAS 222411; collected by Htun Win, Kyi Soe Lwin, Awan Khwi Shein and Hla
Tun, 31 March 2001. CAS 222412—222413 (Field numbers JBS 8240-41); Che Stream, Min Dat
Township, Min Dat District, Chin State (21°21'14.9”N, 93°56’08.3”E, 1298 m.); collected by Htun
Win, Thin Thin, Kyi Soe Lwin and Awan Khwi Shein, 30 March 2001.
ETYMOLOGyY.— The specific epithet is a patronym honoring Carl Gans (born 1923), who has
made substantial contributions to the herpetology of tropical Asia and who has been a strong influ-
ence on my own professional career and that of many other herpetologists and vertebrate morphol-
ogists around the world. The name is masculine and is formed in the genitive case.
DEFINITION.— A moderate sized Cyrtodactylus, snout-vent length to 63 mm; body relatively
slender, limbs and digits relatively short; one pair of enlarged postmental scales, in broad contact
with one another behind mental; 20—25 rows of rounded, conical dorsal tubercles; 36—40 ventral
scales between lowest rows of dorsal tubercles; no discrete ventrolateral folds; shallow precloacal
groove in males, 16—29 large precloacal pores in a single, strongly angled series (pores smaller and
not recessed in females). Seven widened subdigital lamellae beneath basal phalanx of 4' toe of pes,
11 narrow lamellae beneath more distal phalanges of same toe (in holotype). Subcaudal scalation
without enlarged midventral plates. Dorsal pattern of approximately 10 narrow, dark brown trans-
verse bands from occiput to sacrum. Tail with alternating brown and white bands.
DESCRIPTION (based on holotype, CAS 222414).— Adult male, SVL 60.8 mm. Head moder-
ately long (HeadL/SVL ratio 0.28), relatively narrow (HeadW/HeadL ratio 0.58), not depressed
(HeadH/HeadL ratio 0.40), distinct from neck. Lores and interorbital region inflated, canthus ros-
tralis not well developed. Snout short (SnEye/HeadL ratio 0.36); much longer than eye diameter
(OrbD/SnEye ratio 0.66); scales on snout and forehead rounded, granular to weakly conical, inter-
mixed with scattered small tubercles behind level of orbits; scales on snout much larger than those
on occipital region. Eye relatively small (OrbD/HeadL ratio 0.24); pupil vertical with crenelated
margins; supraciliaries short, blunt. Ear opening round, moderately large (EarL/HeadL ratio 0.07);
eye to ear approximately equal to diameter of eyes (EyeEar/OrbD ratio 1.03). Rostral approximate-
ly 75% as deep (1.9 mm) as wide (2.5 mm), divided dorsally by rostral groove; two enlarged
476
FIGURE 7.
Cyrtodactylus gansi, sp. nov.
(CAS 222414) from Min Dat
Township, Chin State, Myanmar.
Note the relatively short digits,
narrow dark crossbands, and lack
of transverse rows of white punc-
tations. Scale bar = 10 mm.
Holotype of
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
supranasals separated by a single, somewhat smaller, anterior inter-
nasal and two much smaller posterior internasals; rostral in contact
with supralabial I, supranasals, and anterior internasal; nostrils oval,
laterally oriented, each in broad contact with rostral and also sur-
rounded by supranasal, first supralabial, and two postnasals; pig-
mented narial flap partially occludes posterior half of nostril; 2—5
rows of scales separate orbit from supralabials. Mental triangular,
wider (2.3 mm) than deep (1.6 mm); one pair of greatly enlarged
postmentals, each approximately 40% size of mental; left and right
postmentals in broad medial contact with no intervening granules,
each member of pair bordered laterally by first infralabial and an
enlarged lateral chinshield, the pair bordered posteriorly by 7 slight-
ly enlarged chin scales. Infralabials bordered by 2—3 rows of enlarged
scales, largest anteriorly and laterally. Throat scales small, rounded,
granular. Supralabials (to midorbital position) 8; enlarged supralabi-
als to angle of jaws 11 (left)—12 (might); infralabials 11; interorbital
scale rows across narrowest point of frontal bone 13.
Body moderately slender, elongate (TrunkL/SVL ratio 0.46)
without discrete ventrolateral folds. Dorsal scales heterogeneous,
mostly rounded to weakly conical granules with pitted or rugose sur-
faces, intermixed with regularly arranged small (3 times granule
size), rounded, conical tubercles extending from occipital region on
to back and base of tail; tubercles on nape smaller than those on body
dorsum; tubercles in approximately 22 longitudinal rows at midbody;
53 tubercles in paravertebral row from occiput to mid sacrum. Ventral
scales much larger than dorsal, cycloid, imbricate to subimbricate;
enlarged between precloacal pores and vent; midbody scale rows
across belly between lowest rows of dorsal tubercles 36; scales on
throat minute, granular, grading into larger scales on chest.
Precloacal pores large, in a single, strongly angled series of 17 lying
in a shallow precloacal groove (Fig. 8); no femoral pores. Scales on
palm and sole smooth, flattened; scales on dorsal aspects of
hindlimbs granular, conical, similar to dorsal scales, with larger, con-
ical tubercles
interspersed.
Dorsal
scales of
proximal forelimbs imbricate, without tuber-
cles; scales of forearms heterogeneous with few
scattered conical tubercles.
Fore- and hindlimbs relatively slender;
forearm (ForeaL/SVL ratio 0.15) and tibia
(CrusL/SVL ratio 0.17) moderate in length;
digits relatively short, strongly inflected at each
joint, all bearing robust, recurved claws: sub-
digital lamellae widened beneath basal phalanx
to approximately half digital width; lamellae
from first proximal scansor greater than twice
FIGURE 8. Cloacal region of holotype of Cyrtodactylus
gansi, sp. nov. (CAS 222414). Note the strongly angled and
shallowly recessed series of large precloacal pores and the
enlarged row of scales posterior to the pore-bearing scales.
Arrow indicates apex of pore-bearing scale series.
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 477
largest palm scale to basalmost digital inflection: 3-4—5—6-3 (manus) and 2—5—6—7-S (pes); lamel-
lae from basalmost digital inflection to toe tip, not including ventral claw sheath: 7-8—10—10—9
(manus) and 8—9—11—11—11 (pes); one to several rows of small, non-lamellar granules between
basal and distal lamellar series; interdigital webbing present but weakly developed. Relative length
of digits (manus; measurements in mm in parentheses): [V (5.1) > III (4.8) > II (4.3) > V (4.0) >I
@>)(pes)s1V (6.2) > V @:8)> Il 6.6) > 4:8) > 1.8):
Partially regenerated tail longer than body (TailL/SVL ratio 1.24), slender, cylindrical in cross
section. Scales arranged in regular segments; dorsal and ventral scales rounded at free margins.
Eight circumferential rows per segment; pygal segments with one transverse row of 12 enlarged
conical tubercles, each tubercle separated from next by 2—5 smaller scales; more distally tubercles
decreasing to 6 then 2 per segment; tubercles absent on distal 3/4 of tail; posteriormost caudal
scales narrow and elongate. Subcaudal scales much larger than dorsal, 3 rows per segment; no
enlarged median plates. Cloacal spurs with 3 enlarged, smooth, flattened, pointed scales on each
side of tail base.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3—4—5-3 for
manus and 2-34-54 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 17.5 postpygal caudal vertebrae in the partly regen-
erated tail in holotype (31—34 in paratypes with complete tails, 1.5 in paratype with broken tail).
One pair of slender, crescentic cloacal bones, with enlarged lateral flanges present in males at level
of second pygal vertebrae (absent in female paratype). Cloacal bones relatively large in all males
except subadult paratype (CAS 222411), which exhibits incomplete ossification of some long
bones (juvenile paratype not examined osteologically). Endolymphatic sacs not enlarged extracra-
nially.
Coloration (in preservative). Base color light to medium brown with dark transverse markings,
irregular from nape to shoulder, forming 7 more-or-less continuous crossbands from forelimb
insertion to sacrum. Bands terminate abruptly on flanks; small, dark spots irregularly positioned at
ventrolateral margins, between ends of crossbands. Top of head medium brown with a diffuse set
of irregular dark brown spots. Canthal region somewhat darker than rest of snout. A brown streak
running from behind orbit to above ear, in conjunction with dark spots on neck forming a broken
nape band. Limbs and feet bearing dark spots. Dark markings on tail forming 10 bands, some part-
ly fused with one another, on original portion of tail. Dark tail bands slightly narrower than lighter
interstices. Tail patterning does not extend on to venter. Venter beige with scattered dark pigment
on all surfaces, darkest under thighs and
around cloaca. Palms and soles of feet
darker than remainder of venter.
Color in life similar to that in preser-
vative but rims of orbit and lateral tuber-
cles yellow, with a yellowish tinge to the
lighter areas of the limbs (Fig. 9).
VARIATION.— Comparative mensural
data for the holotype and paratypes are . :
presented in Table 3. Paratypes similar to Ficure 9. Living paratype specimen of Cyrtodactylus gansi, sp.
holotype in most respects except as noted. _noy. (CAS 222411) illustrating the dorsal coloration of the species.
Scale rows between lowest rows of tuber- Photo by Hla Tun.
cles at midbody 36-40. Rows of dorsal tubercles at midbody 20-25. Precloacal pores in males
16-29, in a continuous series. Pores present, but much smaller in the single adult females paratype
(CAS 222413), a single poreless scale dividing the rightmost pore from a continuous series of 13;
478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
TABLE 3. Mensural data for the types of Cyrtodactylus gansi, sp. nov.
Abbreviations as in Materials and Methods section; all measurements in mm.
|CAS 222414 CAS 222411 CAS 222412 CAS 222413 CAS 226144 CAS 226145
Holotype Paratype Paratype Paratype Paratype Paratype
Sex | male male male female male juvenile
SVL | 60.8 46.5 57.3 62.4 62.3 29.3
ForeaL 8.9 7 8 8.6 8.5 4:5
CrusL eeeet03 7.8 9.5 11 10.5 5.3
TailL (entire) 75.6 53.9 6.4 qf) 54.2 30.4
TailL (portion 28.6 — broken ae 47.8 is
regenerated)
TailW 5.9 Bhs) 5.8 4.2 4.9 Mp)
TrunkL 28.1 20.6 25.7 30.3 252 11.5
HeadL 17.1 12.2 15:5 16.6 7/3} 8.7
HeadW 10 7.8 9.4 9.9 10.2 5.6
HeadH 6.8 5 5.8 6 6.6 3.8
OrbD 4 Dal, 3.9 4 4.3 Dell
EyeEar 4.2 39 4.5 4.6 5 2.6
SnEye 6.1 4.9 5.8 6.6 6.4 3.4
NarEye 4.5 Bul 3.8 4 4.6 Dee
Interorb 5.6 3.4 5 Sie) 5.5 333
EarL 1.2 1.1 1.3 1.4 1.4 0.6
2 Hell
Internar 9) 1.6 17 Dail
no precloacal groove in female specimen, no pores or groove in juvenile (CAS 226145).
Supralabials to midpoint of orbit 7 on right side of CAS 222412 and 226144); infralabials 9 (CAS
226144, right) or 10 (CAS 222412 and CAS 226144, left). Supranasal scales in contact anteriorly
in CAS 222413, one posterior internasal only in CAS 226144, two posterior internasals arranged
longitudinally in CAS 222411.
Color pattern less fragmented on nape and shoulders in some specimens (CAS 222412,
226144), yielding 10 transverse bands to sacrum. Dorsal markings less bold and continuous in
juvenile (CAS 226145). Thirteen (CAS 222411) or 15 (CAS 222413) dark tail bands in specimens
with original tails.
DIAGNOsIS.— Cyrtodactylus gansi may be distinguished from all congeners on the basis of its
possession of short digits, a single, strongly angled and somewhat recessed series of precloacal
pores (16-29) in males (pores smaller and not in groove in females), absence of femoral pores,
20-25 longitudinal rows of dorsal tubercles; 36-40 ventral scales between lowest rows of dorsal
tubercles (no discrete ventrolateral folds), subcaudal scalation without enlarged midventral plates,
and dorsal pattern of approximately 10 narrow, dark transverse bands from occiput to sacrum,
sometimes fragmented anteriorly.
Among other species from Myanmar it is most similar to C. khasiensis and C. ayeyarwadyen-
sis (see account of latter species abive), from which it may be distinguished by its shallow precloa-
cal groove (in males) and its dorsal color pattern.
DISTRIBUTION.— Cyrtodactylus gansi is currently known only from high elevation (750-1300
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 479
m) at the type locality of Che Stream, Min Dat Township, Min Dat District in Chin State, in the
southern Chin Hills (Fig. 3). The recently described C. slowinskii, previously known only from
Alaungdaw Kathapa National Park, in the Sagaing Division, is the only congener thus far found at
this locality (CAS 222415).
Cyrtodactylus wakeorum Bauer, sp. nov.
Figs. 10-11
HoLotypPe.— California Academy of Sciences (CAS) 221935 (Field number JBS 8948), adult
female; Rakhine Yoma Elephant Range, Gwa, Gwa Township, Rakhine State, Myanmar
(17°35’02.4’N, 94°40’44.2”E): collected by Htun Win, Thin Thin, Kyi Soe Lwin and Awan Khwi
Shein, 2 June 2001.
PARATYPE.— CAS 226148 (Field number JBS 8408), juvenile; Ye Bya Stream, Rakhine Yoma
Elephant Range, Gwa Township, Rakhine State, Myanmar (17°41'17.5”N, 94°38’50.8”E, 180 m.);
collected by J. B. Slowinski, GO.U. Wogan, Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi
Shein and Hla Tun, 25 April 2001.
ETYMOLOGY.— The specific epithet honors Marvalee H. Wake (born 1939) and David B.
Wake (born 1936) of the University of California, Berkeley for their contributions to herpetology,
vertebrate morphology and evolutionary biology and for their valuable mentoring of many gradu-
ate students, myself included. It gives me particular pleasure to name and describe this species on
the occasion of the nominal retirement of my former dissertation advisor and mentor, Marvalee
Wake. The specific epithet is masculine (mixed : ee oe
gender) and is formed in the genitive plural.
DEFINITION.— A small sized
Cyrtodactylus, snout-vent length to 64 mm;
body moderately slender, limbs and digits rela-
tively short; one pair of enlarged postmental
scales in broad contact behind mental; dorsum
with relatively smooth texture, with 24 longitu-
dinal rows of small, oval to rounded, keeled
tubercles; 31 ventral scales across midbody; no
precloacal groove, 12 precloacal pores in a sin-
gle series in female holotype, no femoral pores.
Ten subdigital lamellae beneath 4' toe of pes
distal to basal digital inflection, six broad
lamellae basal to inflection. Subcaudal scales
not forming broad transverse plates. Dorsal pat-
tern of 6 thin dark bands (one nuchal and five
between shoulders and sacrum), each bordered
posteriorly by a thin white line. Head without
dorsal pattern. Tail with alternating light and
dark banding.
DESCRIPTION (based on holotype, CAS
221935).— Adult female containing two eggs.
Ficure 10. Holotype of Cyrtodactylus wakeorum, sp.
Snout-vent length 63.8 mm. Head relatively
short (HeadL/SVL ratio 0.25), moderately nar-
row (HeadW/HeadL ratio 0.59), not depressed
(HeadH/HL ratio 0.43), distinct from neck.
noy. (CAS 221935) from Rakhine Yoma Elephant Range.
Rakhine State, Myanmar. Note the relatively short digits,
light-edged dark crossbands, and patternless head dorsum of
this gravid female. Scale bar = 10 mm.
480. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
Lores and interorbital region weakly inflated, canthus rostralis not particularly prominent. Snout
relatively short (SnEye/HeadL ratio 0.39); longer than eye diameter (OrbD/SnEye ratio 0.60);
scales on snout and forehead rounded, granular, flattened to slightly conical, becoming heteroge-
neous posterior to mid-frontal region; scales on snout larger than those on occipital region. Eye rel-
atively large (OrbD/HeadL ratio 0.25); pupil vertical with crenelated margins; supraciliaries short,
without projecting spines. Ear opening oval, small (EarL/HeadL ratio 0.08); eye to ear distance
greater than diameter of eyes (EyeEar/OrbD ratio 1.38). Rostral 64% as deep (1.6 mm) as wide (2.5
mm), incompletely divided dorsally by well developed rostral groove; two enlarged supranasals
separated by a much smaller anterior internasal and two even smaller posterior internasals; rostral
in contact with supralabial I, supranasals, and anterior internasal; nostrils circular, each surround-
ed by supranasal, rostral, first supralabial, one postnasal, and crescentic nasal, rostral contact of
narial border extensive; narial flap partially occludes posterior half of nostril; 1-3 rows of scales
separate orbit from supralabials. Mental subtriangular, wider (2.1 mm) than deep (1.6 mm); one
pair of enlarged postmentals, each 50% size of mental, in broad contact with one another medial-
ly, bordered anterolaterally by first infralabial (and second infralabial on left side only), posterolat-
erally by enlarged lateral chinshield, and posteriorly by 3 chin granules. Infralabials bordered medi-
ally by 1-2 rows of enlarged scales. Supralabials (to midorbital position) 8 (left) —9 (right);
enlarged supralabials to angle of jaws 11; infralabials 10; interorbital scale rows across narrowest
point of frontal 12.
Body relatively slender, relatively short (TrunkL/SVL ratio 0.40) with ventrolateral folds
indistinct. Dorsal scales small, granular, conical, with regularly arranged small tubercles extending
from frontal region on to tail base; each tubercle oval to rounded, bearing a single prominent keel;
tubercles becoming smaller and less prominently keeled on flanks; largest keeled tubercles in
approximately 24 regular rows at midbody. Ventral scales larger than dorsal, smooth, subimbricate
laterally, imbricate across midventer; somewhat larger than elsewhere along ventral midline of
abdomen, and especially in precloacal region; midbody scale rows across belly to edge of flanks
(as demarcated by dorsal color pattern) 31; gular region with homogeneous scalation. Precloacal
pores in a single continuous series of 12; bordered posteriorly by a row of greatly enlarged pore-
less scales. No femoral pores or precloacal groove. Scales on palm and sole smooth, rounded;
scales on dorsal aspects of forelimbs smooth, subimbricating, heterogeneous but without tubercles.
Hindlimbs with small scattered tubercles.
Fore and hindlimbs relatively stout; forearm short (ForeaL/SVL ratio 0.13); tibia short
(CrusL/SVL ratio 0.16); digits moderately short, strongly inflected at basal interphalangeal joints,
all bearing robust, slightly recurved claws; subdigital lamellae rounded, smooth, without scansor-
ial surfaces; lamellae distal to much enlarged scale at basal digital inflection and not including ven-
tral claw sheath: 44—5—5-—5 (manus), 3—4-5—6-5 (pes), proximal 1—3 fragmented on most digits;
enlarged basal lamellae (> twice size of palmar scales) to and including enlarged scale at basal
inflection: 6-7—9—9-8 (manus), 7—7—10—10—11 (pes); interdigital webbing absent. Relative length
of digits (manus; measurements in mm in parentheses): III (4.3) > IV (4.0) > II (3.4) ~ V (3.3) >I
(Qs (pes) 2 1V, 6-9) > Ill 6.2) = V 6.1) > G.9)>1 C9).
Tail partly regenerated, slightly longer than body (TailL/SVL ratio 1.08), slender, tapering;
divided into indistinct segments, each 8—9 dorsal scale rows in length; two rows of enlarged, flat-
tened, keeled tubercles positioned paravertebrally on tail base only, remaining dorsal caudal scales
small, smooth, rectangular; subcaudal scales larger (3 per tail segment), smooth, imbricate not
forming a single series of transverse plates. Two enlarged, smooth, conical postcloacal spurs on
each side of tailbase.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3-4—5-3 for
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 48]
manus and 2—3—4—5 + for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 17.5 post pygal caudal vertebrae in original tail in
holotype (25 post pygal vertebrae in original tail of juvenile paratype). Adult female holotype con-
taining two large eggs. No cloacal bones present in female, not discernable in juvenile. Juvenile
paratype with most bones incompletely ossified. Endolymphatic sacs not enlarged extracranially.
Coloration (in preservative). Base color mid brown. Boldly marked with a series of chocolate
brown bands, each outlined posteriorly by a thin cream to whitish border one scale width in thick-
ness. Occipital band turns at a right angle above and behind ear and extends anteriorly to orbit and
under eye to loreal region, portion on temporal region almost completely surrounded by thin white
border. Dorsum of head unpatterned. Nuchal markings paired, borders just touching one another
along midline. Five additional bands between shoulders and sacrum, each successively more
chevron-shaped, becoming more asymmetrical posteriorly. Dark markings faded on flanks. Limbs
more-or-less uniform mid-brown with scattered small whitish markings. Hindlimb insertions with
a large chocolate brown blotch at anterior border. Venter beige tinged by the light brown speckling
of individual scales, except along midline. Tail with alternating chocolate brown and mid brown to
cream banding. Paler bands wider than darker basally. Tail venter with scattered dark pigmentation,
especially distally. Dorsal caudal color pattern extends onto ventrum, especially posteriorly.
Base color in life pinkish brown, venter
whitish. Light borders around dark dorsal mark-
ings, and supraciliary scales yellowish to yel-
lowish-orange. Iris golden brown (Fig. 11).
VARIATION.— Comparative mensural data
for the holotype and paratype are presented in
Table 4. The juvenile paratype (CAS 226148) is
similar to the holotype in most respects except:
left postmental scale fragmented into two
FiGurE 11. Living holotype of Cyrtodactylus wakeorum,
scales; nostril bordered posteriorly by three
postnasals; enlarged supralabial scales to mid-
dle of eye 9 (left) and 10 (right), to corner of
sp. nov. (CAS 221935) illustrating the yellowish edging of
the dorsal bands and the supraciliary scales. Photo by Hla
Tun.
mouth 11 (left) and 12 (right), infralabials 9 (left) and 10 (right); precloacal pores not developed,
tail tip broken, occipital band not connected to temporal markings, nape markings fused to form a
single band, more posterior bands paired, dark blotches at posterior border of hindlimb insertions.
Both the holotype and paratype exhibit some skin abrasions, suggesting that the skin in life may be
relatively delicate.
DIAGNOsIS.— Cytrodactylus wakeorum may be distinguished from all congeners on the basis
of its possession of relatively short limbs and digits, one pair of enlarged postmental scales in broad
contact behind mental; dorsum with relatively smooth texture, with 24 longitudinal rows of tuber-
cles, 31 ventral scales across midbody, ventrolateral folds absent, no precloacal groove, 12 pre-
cloacal pores in a single series in female holotype, no femoral pores, subcaudal scales not forming
broad transverse plates, dorsal pattern of 6 thin dark bands (one nuchal and five between shoulders
and sacrum), each bordered posteriorly by a thin white line, and head without dorsal pattern.
Cyrtodactylus wakeorum is superficially most similar to C. consobrinoides and, to a lesser
extent, C. annandalei. It may be distinguished from the former by its possession of lack of subcau-
dal transverse plates, and its patternless head dorsum. It differs from the latter in its greater num-
ber of dorsal tubercle rows (24 vs 16-18), smaller number of ventral scale rows (31 vs 43), and
light borders around only the posterior margins of the dark dorsal markings. Unfortunately the
absence of adult male specimens precludes the use of certain precloacal and femoral pore charac-
482 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
ters which might further distinguish C. wakeorum from its congeners. Comparisons with other
species are provided following the description of all new taxa.
DISTRIBUTION.— Cyrtodactylus wakeorum is known only from Rakhine Yoma Elephant
Range, southern Rakhine State (Fig. 3). It is there sympatric with C. ayeyarwadyensis.
Cyrtodactylus russelli Bauer, sp. nov.
Figs. 12-14
HoLoryPe.— California Academy of Sciences (CAS) 226137 (Field number JBS 15525),
adult male; upper Nat E-Su Stream, Htamanthi Wildlife Sanctuary, Hkamti Township, Sagaing
Division, Myanmar (25°28’27.7”N, 95°37'20.5”E, 227 m.); collected by Htun Win, Thin Thin and
Awan Khwi Shein, 8 November 2002.
PARATYPE.— CAS 226140 (Field number JBS 18088), adult female; Hepu Stream, Indawgyi
Wildlife Sanctuary, Mohuyin Township, Kachin State, Myanmar (25°05’38.2”N, 96°22’49.0”E);
collected by J.A. Wilkinson, G.O.U. Wogan, J.V. Vindum, Thin Thin, Kyi Soe Lwin, Awan Khwi
Shein and Hla Tun, 16 May 2003.
ETYMOLOGY.— The specific epithet honors my friend and colleague Anthony P. Russell (born
1947) of the University of Calgary for his many contributions to the study of gekkonid lizards and
for his influence on my professional career. The epithet is masculine and is formed in the genitive
singular. ee ee:
DEFINITION.— A large sized Cyrtodac-
tylus, snout-vent length to 116 mm; body mod-
erately slender, elongate with well developed
ventrolateral folds, limbs stout, digits long; one
pair of enlarged postmental scales in broad con-
tact behind mental; dorsum with 22 longitudinal
rows of small, conical to keeled tubercles;
35-41 ventral scales across midbody to ventro-
lateral folds; no precloacal groove, 15 precloa-
cal pores in a single series in male holotype,
16-19 femoral pores on each thigh separated
from precloacal pores by a diastema (precloacal
and femoral pores absent in female paratype).
Thirteen subdigital lamellae beneath 4 toe of
pes distal to basal digital inflection, nine broad
lamellae basal to inflection. Subcaudal scales
forming broad transverse plates, but distinctly
narrower than tail. Dorsal pattern poorly
defined, including dark elongate blotches or
bands and scattered lighter blotches on flanks.
Tail with alternating light and dark banding.
DESCRIPTION (based on holotype, CAS
2261337, except as noted).— Adult male with
abdominal incision from tissue removal. Snout-
vent length 105.7 mm. Head relatively long
(dead Eis VE ratio 0.28), wide (HeadW/HeadL (CAS 226137) from Htamanthi Wildlife Sanctuary, Sagaing
ratio 0.64), not depressed (HeadH/HL ratio _ pjivision, Myanmar. Note the long digits, robust body, and
0.41), distinct from neck. Lores and interorbital — obscure dorsal pattern. Scale bar = 10 mm.
FiGurE 12. Holotype of Cyrtodactylus russelli, sp. nov.
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 483
region strongly inflated, canthus rostralis well developed. Snout moderately short (SnEye/HeadL
ratio 0.39); longer than eye diameter (OrbD/SnEye ratio 0.60); scales on snout and forehead round-
ed, granular to weakly conical, intermixed with scattered small tubercles posterior to fronto-pari-
etal suture; scales on snout much larger than those on occipital region. Eye relatively small
(OrbD/HeadL ratio 0.23); pupil vertical with crenelated margins; supraciliaries short, each posteri-
or supraciliary bearing a short spines Ear opening rounded, partly occluded by horizontal fold of
skin, small (EarL/HeadL ratio 0.04); eye to ear distance greater than diameter of eyes
(EyeEar/OrbD ratio 1.17). Rostral 64% as deep (2.6 mm) as wide (4.1 mm), divided dorsally by a
weakly developed inverted “V”-shaped rostral groove extending about 20% of scale height; two
enlarged supranasals separated by two somewhat smaller and irregular internasals; rostral in con-
tact with supralabial I, supranasals, and two internasals; nostrils oval, laterally oriented, each sur-
rounded by one supranasal, rostral, first supralabial, one postnasal, and a larger crescentic nasal,
rostral contact of narial border extensive; narial flap partially occludes posterior 2/3 of nostril; 3-5
rows of scales separate orbit from supralabials. Mental subtriangular, much wider (3.7 mm) than
deep (3.1 mm); one pair of enlarged postmentals, each approximately 40% size of mental, in broad
contact with one another medially, bordered anterolaterally by first infralabial, posterolaterally by
enlarged lateral chinshield, and posteriorly by 4 small chin granules. Infralabials bordered medial-
ly by 2—3 rows of enlarged scales. Supralabials (to midorbital position) 9; enlarged supralabials to
angle of jaws 11 (left) to 12 (right); infralabials 10 (left) to 11 (right); interorbital scale rows across
narrowest point of frontal 28.
Body relatively robust, elongate (TrunkL/SVL ratio 0.47) with weakly denticulate, well devel-
oped ventrolateral folds. Dorsal scales heterogeneous, rounded, granular to weakly conical, inter-
mixed with regularly arranged small (6—8 times granule size), keeled, rounded tubercles extending
from parietal region on to back and tail base; tubercles on nape more strongly conical and without
keels; tubercles in approximately 22 longitudinal rows at midbody; 42 tubercles in paravertebral
row from occiput to mid sacrum. Ventral scales much larger than dorsal, cycloid, smooth, subim-
bricate to imbricate; not enlarged under thighs or between precloacal pores and vent; midbody scale
rows across belly between ventrolateral folds 35; scales on throat minute, granular, grading into
larger scales on chest. Precloacal pores in a single continuous series of 15, in a weakly developed
groove; 19 femoral pores on left thigh, 16 on right thigh, each series separated from precloacal pore
series by 3 poreless scales; Scales on palm and sole smooth, flattened; scales on dorsal aspects of
limbs granular to conical, similar to dorsal scales, with larger, conical tubercles interspersed.
Fore and hindlimbs relatively short, stout; forearm short (ForeaL/SVL ratio 0.15); tibia short
(CrusL/SVL ratio 0.17); digits long, strongly inflected at basal interphalangeal joints, all bearing
robust, slightly recurved claws; subdigital lamellae rounded, smooth, without scansorial surfaces,
widened beneath basal phalanx to almost width of toes; lamellae from first proximal scansor
greater than twice largest plam scale to basal digital inflection: 5-7—7—7—8 (manus), 6—8-8—9-8
(pes); lamellae from basal inflection to toe tip, not including ventral claw sheath: 10—10—13—10-11
(manus), 9-11—13—13-13 (pes); one to several rows of distal lamellae fragmented; interdigital
webbing present, especially between digits II and III and III and IV, but weakly developed. Relative
length of digits (manus; measurements in mm in parentheses): IV (10.4) > V (10.0) > II (9.3) >
(8.2) > I (6.0); (pes): V (11.6) ~ IV (11.5) > If (10.9) > I (9.0) > I (6.0).
Tail in holotype broken at base. Original tail in paratype longer than body (TailL/SVL ratio
1.19), slender, tapering, slightly depressed in corss section. Scales arranged in regular whorls; ven-
tral scales rectangular, dorsal scales rounded at free margins or hexagonal. 9 circumferential rows
of dorsal scales per tail segment; basalmost postpygal segment with one transverse row of 10
enlarged keeled tubercles, each tubercle separated from the next by 1-4 smaller scales; more dis-
484 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
tally tubercles decreasing by pairs to just two per transverse row; posteriormost caudal scales nar-
row and relatively elongate. Subcaudal scales much larger, 2 per tail segment, each transversely
enlarged but distinctly narrower than tail; tail segmentation weakly demarcated ventrally. Cloacal
spurs with 34 slightly enlarged, smooth, rounded scales on each side of tailbase.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3-4—5-3 for
manus and 2-34-54 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 1.5 post pygal caudal vertebrae in regenerated tail in
holotype (39 post pygal vertebrae in original tail of paratype). One pair of crescentic cloacal bones
present, expanded posterolaterally (absent in female paratype). Endolymphatic sacs not enlarged
extracranially.
Coloration (in preservative). Dorsum mid- to dark brown with poorly differentiated darker
markings forming elongate paravertebral blotches and more-or-less continuous longitudinal bands
of variable thickness on the dorsolateral margins of trunk (Fig. 12); lighter longitudinally oriented
blotches on flanks. A series of three grayish spots with irregular margins across shoulders. Forelimb
insertions and axillae pale grayish. Forelimbs mottled brown, hindlimbs mottled brown but with
some irregular grayish barring proximally; head darker than trunk, without obvious markings; lore-
als and area beneath orbit somewhat darker than rest of head; labial pale with scattered dark pig-
ment on all scales. Venter grayish cream with dark pigment on limb margins and on throat and mar-
gins of jaws; scattered pigment across all ventral scales. Tail dorsum with alternating wider darker
bands and narrower light brown bands; 9 dark bands on original tail; caudal color pattern extends
onto venter, more distinct posteriorly
(based on paratype).
Pattern much bolder in life, consist-
ing of a series of wavy-edged, transverse
dark markings with thin, pale borders.
Anteriormost marking continuous across " eet “Tn a
occiput, nape and shoulder markings frag- ie "as —
mented, four continuous markings across Ps ee bes RRS ee |
body and sacrum (Fig. 13). Large brown FiGuRE 13. Living paratype of Cyrtodactylus russelli, sp. nov.
patch continuous from snout to anterior of (CAS 226140) illustrating the dorsal pattern of wavy dark cross-
bars. Photo by Hla Tun.
forelimb insertion (Fig. 14). Iris bronze.
VARIATION.— Compara-
tive mensural data for the holo-
type and paratype are present-
ed in Table 4. Adult female
paratype (CAS 226140) differs
from holotype in: internasals 2
but arranged along body axis,
not transversely; rostral crease
Straight, not “V”-shaped; 9
(right) to 10 (left) supralabials
to middle of eye, 11 (right) to
12 (left) to corner of mouth, 9
(right) to 10 (left) infralabials;
41 ventral scales between ven-
trolateral folds; 44 tubercles in
FiGurE 14. Lateral view of living paratype of Cyrtodactylus russelli, sp. nov.
(CAS 226140) illustrating the large brown patch on the side of head and neck,
whitish outline of dorsal markings, beige venter, and golden iris. Photo by Hla
paravertebral row from occiput Tun.
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 485
TABLE 4. Mensural data for the types of four new species of Cyrtodactylus.
Abbreviations as in Materials and Methods section; all measurements in mm.
C. wakeorum
C. russelli | C. chrysopylos | C. aequalis
CAS 221935 CAS 226148 | CAS 226137 CAS 226140| CAS 226141 | CAS 222185
Holotype Paratype Holotype Paratype Holotype _ Holotype
Sex female ? Guvenile) male female female female
SVL 63.8 33 105.7 116 79.1 | 90.1
ForeaL 8.5 4.9 16 17.9 | 13.2 | 14
CrusL 10 5.6 18.3 20.5 16.1 oem Ge?
TailL (entire) 69 31.8 ll 138.6 6.2 he OP
TailL (portion 24 bBroken broken _ | broken 49.5
regenerated) |
TailW Sal 2.4 10.6 10.6 | a | 7
TrunkL 28.4 14.1 49.4 52.3 | 36 38.8
HeadL 15.9 10.2 29.2 314 | 22.4 i og
HeadW 9.4 SS 18.6 22.2 | 14 15.9
HeadH 6.9 Soll 11.8 12.9 | 9.3 10.1
OrbD 4 eT 6.8 7.4 6.3 6.6
EyeEar 5.4 Deaf 8 9.3 5.5 6.5
SnEye 6.4 4.1 11.2 Be | 9.4 9.8
NarEye 4.8 2.8 8.2 8.9 | 6.6 | vs
Interorb 5.9 35) 10.6 I leg/ 8.3 | 8.3
EarL 1D 0.8 ] 1.6 | 2.1 2
Internar 1.6 le 3.8 3.6 | 2.6 2)
to mid-sacrum; no precloacal or femoral pores; tail original; dorsal markings more pronounced
than in holotype, consisting of dark brown and grayish mottling; dark paravertebral markings on
nape prominent; dark occipital band present; a narrow whitish line across occiput, fading on tem-
poral region.
DIAGNOSIS.— Cyrtodactylus russelli may be distinguished from all congeners on the basis of
its very large size (to 116 mm SVL), ventrolateral folds well developed, digits long, dorsum with
22 longitudinal rows of tubercles, 35-41 ventral scales across midbody to ventrolateral folds, no
precloacal groove, 15 precloacal pores in a single series in male holotype, 16—19 femoral pores on
each thigh separated from precloacal pores by a diastema (precloacal and femoral pores absent in
female paratype), subcaudal scales forming broad transverse plates, but distinctly narrower than
tail width, and dorsal pattern poorly defined, including dark elongate blotches or bands and scat-
tered lighter blotches on flanks.
Among its congeners in Myanmar, the new species is approached in size only by C. slowinskii
(maximum SVL 108 mm), to which it appears closely related. It differs from this species in its less
well demarcated dorsal pattern, white (vs yellowish) light markings, predominantly transverse
wavy dorsal markings (vs paired blotches), and larger number of precloacal pores (15 vs 9-11) and
femoral pores (16-19 per thigh vs 11-12). Comparisons with other species are provided following
the new species descriptions.
DISTRIBUTION.— Cyrtodactylus russelli is known from two localities (Htamanthi Wildlife
486 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
Sanctuary, Sagaing Division and Indawgyi Wildlife Sanctuary, Kachin State), approximately 85 km
apart in northern Myanmar. The region lies between the border ranges with Assam, India and the
Mangin and Kumon Ranges in the east. It occurs with C. khasiensis at Htamanthi (CAS 226138).
Given the limited sampling in this region of Myanmar, it is impossible to estimate the extent of the
species’ range as a whole. Although no similar geckos have been recorded from Assam, sampling
in this region of India remains inadequate and it may yet be recorded outside of Myanmar.
Cyrtodactylus chrysopylos Bauer, sp. nov.
Figs. 15-17
HOoLoryPe.— California Academy of Sciences (CAS) 226141 (Field number JBS 13417),
adult male; Panlaung-Pyadalin Cave Wildlife Sanctuary, Ywa Ngan Township, Shan State,
Myanmar (21°07’58.4”N, 96°20’25.0’E, 319 m.); collected by GO.U. Wogan, R.S. Lucas, J.V.
Vindum, Htun Win, Thin Thin, Awan Khwi Shein and H. Tun, 14 July 2002.
ETYMOLOGyY.— The specific epithet is derived from the Greek chrysos (golden) and pylos
(gate) and refers to Golden Gate Park, San Francisco the location of the California Academy of
Sciences, which spearheaded the herpetological surveys of
Myanmar that revealed this and many other new taxa. The
name commemorates the 150‘ anniversary of the Academy
and its long history of herpetological research in Asia. I par-
ticularly express my gratitude to the curators and curatorial
staff of the Department of Herpetology for their support of
my research and field expeditions over the past 20 years.
The epithet is a masculine noun in apposition.
DEFINITION.— A moderately sized Cyrtodactylus,
snout-vent length of unique holotype 79 mm; body slender,
elongate, limbs and digits long; one pair of enlarged post-
mental scales in broad contact behind mental; dorsum with
16 longitudinal rows of small, keeled tubercles; 37 ventral
scales across midbody to distinct ventrolateral folds; no pre-
cloacal groove or femoral pores, 10 precloacal pores in a sin-
gle series; a single, much larger, pored scale posterior to pre-
cloacal series and separated from it by a single, enlarged
scale without pores. Thirteen subdigital lamellae beneath 4%
toe of pes distal to basal digital inflection, six broad lamel-
lae basal to inflection. Dorsal pattern of distinct alternating
brown and white bands (one on occiput, one across shoul-
ders, six between limb insertions, one on sacrum, two on
pygal portion of tail).
DESCRIPTION (based on holotype, CAS 226141).—
Adult female with midventral incision from tissue removal.
Snout-vent length 79.1 mm. Head relatively long
(HeadL/SVL ratio 0.28), moderately wide (HeadW/HeadL FIGURE 15. Holotype of (Gye neme
ratio 0.62), not depressed (HeadH/HL ratio 0.42), distinct ae yantios. =. nov. (GAS 226141) poe
from neck. Lores and interorbital region inflated, canthus Panlaung-Pyadalin Cave Wildlife Sanctuary,
rostralis not particularly prominent. Snout moderately long Shan State, Myanmar. Note the long digits,
(SnEye/HeadL ratio 0.42); much longer than eye diameter _ Slender limbs, and alternating light and dark
(OrbD/SnEye ratio 0.67); scales on snout and forehead dorsal nla. Se Oe
N
A
Sil
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 487
rounded, granular, flattened to slightly conical, becoming heterogeneous posterior to frontal region;
scales on snout larger than those on occipital region. Eye large (OrbD/HeadL ratio 0.28); pupil ver-
tical with crenelated margins; supraciliaries short, with small, blunt spines. Ear opening oval, large
(EarL/HeadL ratio 0.09); eye to ear distance less than diameter of eyes (EyeEar/OrbD ratio 0.88).
Rostral 56% as deep (1.9 mm) as wide (3.4 mm), dorsal half incompletely divided by rostral
groove; two enlarged supranasals separated by a single, smaller internasals; rostral in contact with
supralabial I, supranasals, and internasal; nostrils circular, each surrounded by supranasal, rostral,
first supralabial, and two postnasals, rostral and supralabial contact of narial border extensive; nar-
ial flap partially occludes posterior third of nostril; 2-4 rows of scales separate orbit from supral-
abials. Mental subtriangular, much wider (3.6 mm) than deep (2.8 mm); one pair of enlarged post-
mentals, each 30% size of mental, in broad contact with one another medially, bordered anterolat-
erally by first infralabial, posterolaterally by enlarged lateral chinshield, and posteriorly by 3 chin
granules. Infralabials bordered medially by 1—3 rows of enlarged scales, largest anterior and later-
al. Supralabials (to midorbital position) 8 (left) —9 (right); enlarged supralabials to angle of jaws
11; infralabials 10; interorbital scale rows across narrowest point of frontal 19.
Body relatively slender, elongate (TrunkL/SVL ratio 0.45); ventrolateral folds small but dis-
tinct, without denticulate margins. Dorsal scales small, granular to weakly conical, with regularly
arranged small tubercles extending from frontal and temporal regions on to tail base; each tubercle
rounded, bearing a single prominent keel; tubercles becoming smaller and less prominently keeled
on flanks; largest keeled tubercles in approximately 16 regular rows at midbody. Ventral scales
much larger than dorsal, smooth, Sten ueate on mabey, imbricate across chest; somewhat aus
er than elsewhere along ventral zi
midline of chest and abdomen,
and especially in precloacal
region; midbody scale rows
across belly between ventrolat-
eral folds 37; gular region with
homogeneous scalation. Pre-
cloacal pores in a single con-
tinuous series of 10 enlarged
scales, bordered posteriorly by
an enlarged poreless median
scale, this in turn bordered pos-
teriorly by an even larger pored
scale (Fig. 16). No femoral
pores or precloacal groove.
Scales on palm and sole FIGURE 16. Cloacal region of Cyrtodactylus chrysopylos, sp. nov. (CAS
smooth, rounded; scales on 226141) showing the precloacal pore-bearing scales, including the very large
dorsal aspects of proximal scale posterior to the main series (arrow). Note also the absence of enlarged
forelimbs smooth to weakly femoral scales.
conical, relatively homogeneous. Hindlimbs and distal forelimbs with scattered conical tubercles.
Fore- and hindlimbs long, stout; forearm long (ForeaL/SVL ratio 0.17); tibia long (CrusL/SVL
ratio 0.20); digits relatively long, strongly inflected at basal interphalangeal joints, all bearing
robust, slightly recurved claws; subdigital lamellae rounded, smooth, without scansorial surfaces,
widened beneath basal phalanx to approximately *4 width of toes; lamellae from first proximal
scansor greater than twice largest plam scale to basal digital inflection: 3—6—5—6-—5 (manus),
5—6-6-6-6 (pes); lamellae from basal inflection to toe tip, not including ventral claw sheath:
488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
10-10—11—12-12 (manus), 11—12—13—13-—14 (pes); one to several rows of distal lamellae fragment-
ed; very weakly developed interdigital webbing present, especially between digits I and III and II
and IV, but weakly developed. Relative length of digits (manus; measurements in mm in parenthe-
ses): IV (7.9) > III (7.2) > V (6.70) > I (6.2) > I (4.7); (pes): IV (9.3) > HII (8.5) > V (8.2) > II (6.7)
> 14:4):
Tail broken at base; 3 enlarged, smooth, blunt, conical postcloacal spurs on each side of tail-
base.
Mensural data are presented in Table 4.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3-4—5-3 for
manus and 2—3—4—5-4 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 0.5 post pygal caudal vertebrae in regenerated tail in
unique holotype. No cloacal bones present in female type. Endolymphatic sacs not enlarged
extracranially.
Coloration (in preservative). Base color a mottled mid brown. Strongly marked with alternat-
ing chocolate brown and white bands. Dark occipital band extending anteriorly to orbit and under
eye to nostril, becoming less well defined anterior of orbit; bordered above by a broken white line
extending to posterior supraciliaries, bordered posteriorly by a thick white line passing through ear
and onto supralabials. Supraciliaries white, a diffuse white line from anterior of orbit to nostril.
Additional pairs of light and dark alternating bands across shoulders, 6 such pairs between limb
insertions and one across sacrum; 2 additional such pairs on pygal portion of tail. Pattern roughly
bilaterally symmetrical, faded on lower flanks. Dorsum of head with 4 diffuse dark marks on pari-
etal table, largest at posterior margin of orbit. Posterior supralabials white, anterior supralabials
whith scattered white spots. Forelimbs mottled, with white blotches on limbs and limb insertions;
hindlimbs with ill-defined alternating dark brown and white markings. Hindlimb insertions with
large chocolate brown blotches at posterior border of thighs. Venter beige tinged by the light brown
speckling of individual scales, especially on the borders of the jaws.
Base color in life purplish brown. Larger light dorsal markings with a yellowish cast, smaller
lateral spots whitish. Throat white with a pinkish suffusion. Venter beige to light brown (Fig. 17).
DIAGNOSIS.— Cyrtodactylus chrysopylos may be distinguished from its congeners by its rela-
tively long digits, 16 longitudinal rows of dor-
sal tubercles; 37 ventral scales across midbody
to distinct ventrolateral folds, 10 precloacal
pores in a single series, a single, greatly
enlarged pored scale posterior to apex of pre-
cloacal series (Fig. 16), and dorsal pattern of
distinct alternating brown and white bands (one
on occiput, one across shoulders, six between
limb insertions, one on sacrum, two on pygal
portion of tail). FIGURE 17. Living holotype of Cyrtodactylus chrysopy-
As the species is known only froma single /os, sp. nov. (CAS 226141) illustrating the yellowish and
female. the reliability of precloacal and femoral ™ hitish dorsal and lateral markings and the long slender dig-
Sif pete : its. Photo by Hla Tun.
pore characteristics may be called into ques- "
tion, as such features often differ between genders. However, female Cyrtodactylus, if different
from males, typically have a reduced number of pores, or may lack femoral or both femoral and
precloacal pores all together. In this instance the presence in a female of a distinctive large pored
scale posterior to the precloacal series is very likely to be present in males as well. This feature is
unique among Cyrtodactylus and is alone sufficient to diagnose C. chrysopylos from all of its con-
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 489
geners. In addition, its distinctive dorsal pattern of nine bands between occiput and sacrum is
unique.
DISTRIBUTION.— Cyrtodactylus chrysopylos 1s known only from Panlaung-Pyadalin Cave
Wildlife Sanctuary in Shan State, Myanmar at an elevation of 319 m. This locality lies in the west-
ern portion of the extensive hill region occupying east central Myanmar. The new species is sym-
patric with C. peguensis at Panlaung-Pyadalin Wildlife Sanctuary (CAS 226142—226143).
Cyrtodactylus aequalis Bauer, sp. nov.
Figs. 18-20
HoLoryPe.— California Academy of Sciences (CAS) 222185 (Field number JBS 10347),
adult male; Kyaik-Hti-Yo Wildlife Sanctuary, Kyaik Hto Township, Mon State, Myanmar
(17°26’38.1”N, 97°05’56.8”E); collected by Htun Win, Thin Thin and Awan Kwi Shein, 21
November 2001.
ETYMOLOGY.— The epithet is derived from the Latin aequalis meaning “same,” in reference
to the fact that the number of dorsal tubercle rows equals the number of ventral scale rows across
midbody between the ventrolateral folds. This condition is unique among species of Cyrtodactylus.
The epithet is an adjective in the nominative singular.
DEFINITION.— A moderately sized Cyrtodactylus,
snout-vent length of unique holotype 90 mm; body slender,
limbs robust, digits long; one pair of enlarged postmental
scales in broad contact with one another behind mental; dor-
sum with 24 longitudinal rows of relatively large, strongly
keeled tubercles; 24 enlarged ventral scales between distinct
ventrolateral folds; no precloacal groove, 9 minute precloa-
cal pores in female type, 3—4 minute femoral pores separat-
ed from precloacal pores by a diastema. Fourteen subdigital
lamellae beneath 4" toe of pes distal to basal digital inflec-
tion, eight broad lamellae proximal to inflection. Subcaudal
scales forming transverse plates approximately 2/3 width of
tail. Dorsal pattern of paired dark markings bordered by thin
white lines (one on occiput, one across shoulders, five
between limb insertions). Top of head with white vermiform
marks on parietal table and frontonasal region. Tail with
alternating light and dark bands.
DESCRIPTION (based on holotype, CAS 222185).—
Adult female. Snout-vent length 90.1 mm. Head relatively
long (HeadL/SVL ratio 0.27), wide (HeadW/HeadL ratio
0.66), not depressed (HeadH/HL ratio 0.42), distinct from
neck. Lores and interorbital region inflated, canthus rostralis
not especially well developed. Snout moderately long
(SnEye/HeadL ratio 0.41); much longer than eye diameter
(OrbD/SnEye ratio 0.66); scales on snout and forehead FIGURE 18. Holotype of Cyrtodactylus
: : ‘ : : aequalis, sp. nov. (CAS 222185) from
rounded, granular to slightly conical, intermixed with scat- xy aik-Hti-Yo Wildlife Sanctuary, Mon
tered small tubercles posterior to border of orbit; scales on — State, Myanmar. Note the long digits and
snout larger than those on occipital region. Eye relatively bold dorsal and head markings. Scale bar =
large (OrbD/HeadL ratio 0.27); pupil vertical with crenelat- !0™™-
490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
ed margins; supraciliaries short, posterior scales bearing small spines. Ear opening oval, vertically
oriented, relatively large (EarL/HeadL ratio 0.08); eye to ear distance similar to diameter of eyes
(EyeEar/OrbD ratio 0.98). Rostral 56% as deep (1.9 mm) as wide (3.4 mm), no rostral groove; two
enlarged supranasals in broad median contact, a roughlt pentagonal internasal positioned between
rostral and supranasals; rostral in contact with supralabial I, supranasals, and internasal; nostrils
oval, laterally oriented, each in broad contact with rostral and first supralabial, also contacted by
supranasal and two postnasals; pigmented narial flap partially occludes posterior half of nostril;
34 rows of scales separate orbit from supralabials. Mental subtriangular, slightly wider (2.9 mm)
than deep (2.8 mm); one pair of enlarged postmentals, each 60-70% size of mental, in broad con-
tact with one another medially, bordered anterolaterally by first infralabial, posterolaterally by
enlarged lateral chinshield, the pair bordered posteriorly by 8 chin scales including several
enlarged lateral chin scales. Infralabials bordered medially by 2-3 rows of enlarged scales.
Supralabials (to midorbital position) 7; enlarged supralabials to angle of jaws 9 (left) to 10 (right);
infralabials 10 (left) to 11 (right); interorbital scale rows across narrowest point of frontal 19.
Body relatively robust, relatively elongate (TrunkL/SVL ratio 0.43) with very weakly dentic-
ulate ventrolateral folds. Dorsal scales heterogeneous, mostly rounded to weakly conical granules
with pitted or rugose surfaces, intermixed with regularly arranged, moderately large (8-10 times
granule size), strongly keeled to mucronate, rounded tubercles extending from posterior border of
orbits and temporal region on to tail base; tubercles on nape, head, and lower flanks more strong-
ly conical and without keels; tubercles in approximately 24 longitudinal rows at midbody (Fig. 19);
39 tubercles in paravertebral row from occiput to mid-sacrum. Ventral scales much larger than dor-
sal, smooth, imbricate, enlarged along midventral line and in precloacal region; midbody scale
rows across belly to ventrolateral folds 24 (Fig. 20); scales on throat minute, granular, rapidly grad-
ing into much larger scales on chest. Minute precloacal pores in a single continuous series of 9; 3
minute femoral pores on left thigh separated from precloacal pores by diastema of 7 poreless
scales; right thigh with 4 femoral pores (with one poreless scale between the two most distal pores);
femoral pores pierced in row of enlarged femoral scales; no precloacal groove. Scales on palm and
sole smooth, flattened; scales on dorsal aspects of hindlimbs and forearms granular to weakly con-
ical, similar to dorsal scales, with larger keeled tubercles interspersed; dorsal scales of proximal
forelimbs without tubercles.
Fore- and hindlimbs relatively stout; forearm moderate (ForeaL/SVL ratio 0.15); tibia relative-
ly long (CrusL/SVL ratio 0.18); digits long, strongly inflected at basal interphalangeal joints, all
bearing robust, slightly recurved claws; subdigital lamellae rounded, smooth, without scansorial
surfaces, widened beneath basal phalanx to approximately 34 width of toes; lamellae from first
proximal scansor greater than twice largest plam scale to basal digital inflection: 5-6—7—7—7
(manus), 6-8—8—8—8 (pes); lamellae from basal inflection to toe tip, not including ventral claw
sheath: 10—11—13—13-—12 (manus), 11—11—13—14~—16 (pes); interdigital webbing present, especial-
ly between digits If and HI and III and IV, but weakly developed. Relative length of digits (manus;
measurements in mm in parentheses): III (8.8) > IV (8.4) > V (7.2) > II (7.0) > I (5.5); (pes): IV
(11.0) > V (10.4) > III (9.9) > II (8.1) > I (5.8).
Tail partly regenerated, slightly longer than body (TailL/SVL ratio 1.02), slender, tapering,
somewhat depressed in cross section. Scales arranged in regular whorls, 8 dorsal scale rows per tail
segment; pygal segment with one transverse row of 10 enlarged keeled tubercles, each tubercle
separated from next by 1-3 smaller scales; on posterior portion of tail enlarged tubercles reduced
to 4 per transverse row, becoming flattened and eventually unkeeled; tubercles absent on regener-
ated portion of tail, regenerated dorsal caudal scales somewhat irregular in shape, narrow and elon-
gate. Subcaudal scales larger, 2 rows per tail segment, segments not strongly demarcated ventral-
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 491
a
Ficure 19 (left). Dorsum of holotype of Cyrtodactylus aequalis, sp. nov. (CAS 222185) illustrating the large, keeled
tubercles of the dorsum (note: not all tubercle rows are visible in this view).
FIGURE 20 (right). Venter of holotype of Cyrtodactylus aequalis, sp. nov. (CAS 222185) illustrating the large, imbricate
ventral scales and the well-demarcated ventrolateral margin.
ly; median subcaudal scales approximately 2/3 width of tail, forming a row of enlarged plates.
Cloacal spurs with 34 enlarged, smooth, rounded scales on each side of tail base.
Mensural data are presented in Table 4.
Osteology. Parietal bones paired. Stapes imperforate. Phalangeal formulae 2—3—4--5-3 for
manus and 2-34-54 for pes. Presacral vertebrae 26, including 3 anterior cervical (without ribs),
1 lumbar, and 2 sacral vertebrae; 5 pygal and 12.5 post pygal caudal vertebrae in regenerated tail
of unique holotype. No cloacal bones present in female type. Endolymphatic sacs not enlarged
extracranially.
Coloration (in preservative). Base color a medium brown with a series of pairs of dark brown
markings bordered by thin (one scale row wide) white lines. Occipital band continuous with tem-
poral stripe that continues to orbit and passes beneath eye. Ventral white line beneath dark tempo-
ral stripe continues onto supralabials and infralabials; dorsal white line above temporal stripe con-
tinues onto supraciliaries. A series of 5 partly coalescent blotches across nape, a pair of well demar-
cated blotches over shoulders, 5 pairs of increasingly asymmetrical marks between limb insertions,
each with some degree of coalescence with its mate.
Top of head medium brown with a diffuse set of white vermiform marks on parietal table and
frontonasal region. Canthal region crossed by a diffuse whitish stripe. A small white mark at
anteroventral margin of orbit. Labials white with scattered brown blotches. Limbs barred with
white bands extending onto digits, diffuse proximally, more well defined on forearm and crus.
More clearly marked on hindlimb than forelimb. Tail with alternating light and wider dark brown
bands. Asymmetrical distally, with white bands having dark brown centers. Regenerated portion of
tail mottled mid-brown. Venter beige with much scattered brown pigment, especially on posterior
abdomen, limb margins, throat and anterior chest. Underside of tail mottled dark brown with small
white blotches.
DIAGNOSIS.— Cyrtodactylus aequalis may be distinguished from all congeners on the basis of
492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
its long digits, dorsum with 24 longitudinal rows of relatively large, strongly keeled tubercles (Fig.
19), 24 enlarged ventral scales between distinct ventrolateral folds (Fig. 20), no precloacal groove,
9 minute precloacal pores in female type, 3-4 minute femoral pores separated from precloacal
pores by a diastema, subcaudal scales forming transverse plates approximately 2/3 width of tail,
dorsal pattern of paired dark markings bordered by thin white lines (one on occiput, one across
shoulders, five between limb insertions), and top of head with white vermiform markings.
Although the comparison of the precloacal and femoral pore characters of the female type of
C. aequalis with those of males has some limitations, it may be assumed that males also possess
femoral pores, though the number of such pores may be higher. Pore characters aside, the relative-
ly large number of dorsal tubercle rows and small number of ventral scale rows results in equal
counts for these two parameters. This situation is unique in the genus and serves to diagnose the
species from all other congeners. Comparisons with other species are presented below.
DISTRIBUTION.— Cyrtodactylus aequalis is known only from Kyaik-Hti-Yo Wildlife
Sanctuary in Mon State in southern Myanmar, just to the west of the Gulf of Martaban (Gulf of
Mottama).
SPECIES COMPARISONS
The condition of precloacal and femoral scales and pores in males has traditionally been wide-
ly used to distinguish members of the genus Cyrtodactylus (e.g., Smith 1935; Darevsky and
Szezerbak 1997; Bauer 2002). Unfortunately, three of the new species are represented only by adult
females or juveniles. Nonetheless, two of these may be easily distinguished on the basis of unique
features occurring in no other Cyrtodactylus: C. chrysopylos possesses a single, much larger, pored
scale posterior to precloacal series and separated from it by a single, enlarged scale without pores
(Fig. 13) and C. aequalis has greatly enlarged ventral scales (Fig. 17), resulting in an equal num-
ber of ventral midbody scale rows and longitudinal dorsal tubercle rows.
All of the remaining new species from Myanmar lack a deep precloacal groove (a shallow
groove is present in male C. gansi) and can thus be distinguished from C. annulatus (Taylor, 1915),
C. cavernicolus Inger and King, 1961, C. fumosus (Miiller, 1895), C. marmoratus (Gray, 1831), C.
papuensis (Brongersma, 1934), C. philippinicus (Steindachner, 1867), C. pubisulcus Inger, 1958,
C. pulchellus Gray, 1827, C. rubidus (Blyth, 1860), and C. sadleiri Wells and Wellington, 1984;
they may be separated from C. biordinis Brown and McCoy, 1980 by the presence of a single vs.
double row of femoral pores and from the following species by the presence of precloacal pores (at
least in males): C. jellesmae (Boulenger, 1897), C. laevigatus Darevsky, 1964, C. paradoxus
(Darevsky and Szczerbak, 1997), C. sermowaiensis (de Rooij, 1915), and most members of the
subgenus Geckoella (C. albofasciatus [Boulenger, 1885], C. collegalensis [Beddome, 1870], C.
deccanensis |Giinther, 1864], C. jeyporensis |Beddome, 1877], C. nebulosus |[Beddome, 1870], and
C. yakhuna (Deraniyagala, 1945]). All of the new species also lack the acutely angled, short pre-
cloacal pore series (maximum 12 pores) characteristic of C. ingeri Hikida, 1990 and C. yoshii
Hikida, 1990, the denticulate tail margin of C. brevipalmatus (Smith, 1923), and the very short dig-
its of C. brevidactylus Bauer, 2002 and C. (G.) triedrus (Giinther, 1864).
Cyrtodactylus annandalei and C. russelli differ from the following species by the presence of
femoral pores (at least in males): C. adleri Das, 1997, C. angularis (Smith, 1921), C.
ayeyarwadyensis Bauer, 2003, C. condorensis (Smith, 1920), C. consobrinoides (Annandale,
1905), C. elok Dring, 1979, C. fraenatus (Giinther, 1864), C. gansi Bauer, 2003, C. intermedius
(Smith, 1917), C. irianjayaensis Résler, 2001, C. irregularis (Smith, 1921), C. khasiensis (Jerdon,
1870), C. lateralis (Werner, 1896), C. malayanus (de Rooij, 1915), C. matsuii Hikida, 1990, C. old-
hami (Theobald, 1876), C. peguensis (Boulenger, 1893), C. quadrivirgatus Taylor, 1962, C. sumon-
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 493
thai Bauer, Pauwels and Chanhome, 2002, and C. sworderi (Smith, 1925); and from the following
species by the presence of a diastema between precloacal and femoral pore-bearing scales in males:
C. abrae Wells, 2002: C. feae (Boulenger, 1893), C. jarujini Ulber, 1993, C. loriae (Boulenger,
1898), C. louisiadensis (de Vis, 1892), C. malcolmsmithi (Constable, 1949), C. novaeguineae
(Schlegel, 1844), C. papilionoides Ulber and Grossmann, 1991, C. tiomanensis Das and Lim, 2000,
C. tuberculatus (Lucas and Frost, 1900), C. variegatus (Blyth, 1859), and a new species from
Saraburi Province, Thailand (Bauer et al., in press).
Cyrtodactylus russelli differs from the following species in having < 35 ventral scale rows (vs
> 40, except C. darmandvillei, > 36): C. annandalei, Bauer, 2003, C. baluensis (Mocquard, 1890),
C. consobrinus (Peters, 1871), C. darmandvillei (Weber, 1890), C. derongo Brown and Parker,
1973, C. interdigitalis Ulber, 1993, and C. mimikanus (Boulenger, 1914), and from the following
in having 15 precloacal pores (vs. < 11): C. aaroni Giinther and Rosler, 2003, C. agusanensis
(Taylor, 1915), C. gubernatoris (Annandale, 1913), C. redimiculus King, 1962, C. slowinskii
Bauer, 2002, C. wetariensis (Dunn, 1927) and an new species from Kanchanaburi Province,
Thailand (Bauer et al., in press). Cyrtodactylus annandalei differs from these same species (except
C. gubernatoris) by its much smaller adult size (55 mm for an adult female vs. 70 mm and above).
It differs from C. gubernatoris by is greater number of ventral scales across midbody (43 vs. 33).
Cyrtodactylus ayeyarwadyensis, C. gansi and C. wakeorum may be distinguished from the fol-
lowing species by their lack of transversely enlarged subcaudal plates: C. abrae Wells, 2002, C.
baluensis, C. condorensis, C. fraenatus, C. interdigitalis, C. intermedius, C. lousiadensis, C.
sumonthai, C. tuberculatus, C. variegatus, and a new species from Saraburi Province, Thailand
(Bauer et al., in press); from the following by their smaller adult size (<< 79 mm SVL for C.
ayeyarwadyensis, < 64 mm for C. wakeorum, and < 63 mm for C. gansi vs > 90 mm SVL): C. agu-
sanensis, C. angularis, C. consobrinus, C. derongo, C. irianjayaensis, C. jarujini, C. loriae, C.
matsuli, C. mimikanus, C. novaeguineae, C. papilionoides, C. russelli, and C. slowinskii; from C.
adleri, C. elok, C. irregularis, C. oldhami, C. peguensis, C. quadrivirgatus, and C. sworderi in hav-
ing more than 8 precloacal pores; from C. lateralis and C. malayanus by their lower number of ven-
tral scales (< 40 vs > 50); and from C. malcolmsmithi by their greater number of dorsal tubercle
rows (20 or more vs 16).
Cyrtodactylus ayeyarwadyensis and C. gansi differ from the following species in having pre-
cloacal pores only (barely extending onto thighs in some C. ayeyarwadyensis): C. aaroni, C.
annandalei, C. darmandvillei, C. feae, C. gubernatoris, C. redimiculus, C. tiomanensis, C.
wetariensis, and a new species from Kanchanaburi Province, Thailand (Bauer et al., in press).
Female Cyrtodactylus wakeorum may be similarly distinguished from these species, but the male
condition is unknown. Regardless, C. wakeorum differs from all of these species in its particular
dorsal pattern of dark, narrow crossbands edged posteriorly with yellow. Cyrtodactylus
ayeyarwadyensis and C. gansi may be distinguished from C. wakeorum and C. consobrinoides by
the absence of enlarged femoral scales. Differences among C. khasiensis, C. ayeyarwadyensis, and
C. gansi, and between C. wakeorum and C. consobrinoides, respectively, are discussed in the diag-
nostic sections of the new species accounts above.
DISCUSSION
Herpetofaunal diversity of Myanmar has long been underestimated, chiefly owing to limited
collecting activity (Inger, 1999; Slowinski and Wiister, 2000). In the genus Cyrtodactylus this has
been exacerbated by the gross similarity in color pattern of nearly all species (mid-brown with
darker blotches or bands, often with whitish borders) which has made identifications difficult for
494 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
non-specialists. Further, some of the species endemic to the region are known only from a few
specimens, or have been plagued by confusion in the literature. A case in point is Cyrtodactylus
feae. This species was described by Boulenger (1893) from Puepoli, in the Karen Hills of south-
eastern Burma based on one male specimen collected by Leonardo Fea in 1886. Annandale (1905a)
considered three female specimens from “Sinkip Island, East Sumatra” to be referable to the
species, but subsequently (Annandale 1913) reidentified these specimens as juvenile C. consobri-
nus (see also de Rooij 1915). Smith (1935) reexamined the type and concluded that it was a female,
not a male, and determined that Boulenger’s (1893) count of 32 continuous precloacal-femoral
pores was incorrect. Instead Smith regarded the enlarged precloacal and femoral scales to be mere-
ly pitted, not perforated by pores. Constable (1949) followed Smith’s interpretation and referred a
specimen from Calcutta, India to C. feae. However, both this specimen’s provenance and its sup-
posed resemblance to C. intermedius strongly suggest that this specimen is not conspecific with
that described and figured by Boulenger (1893). No additional specimens that are unambiguously
referable to C. feae have since been collected.
Similar confusion regarding precloacal and femoral pore condition is rampant in
Cyrtodactylus. My examination of several hundred specimens suggests that some early authors,
working with hand lenses or inferior low magnification microscopes with poor lighting, may have
overlooked tiny pores and thus mischaracterized certain taxa. Other features that have been wide-
ly used in differentiating members of the genus, including postmental scale condition and the state
of subcaudal scales, while easily observed, appear to have been described differently enough by
different authors as to render literature accounts unreliable for comparative purposes. To assist 1n
the identification of future collections from Myanmar, I provide the following key to the 16 species
thus far recorded from the country.
Key to the Species of Cyrtodactylus Occurring in Myanmar
la Deep iprecloacali(pubic)/sulcus present 225...) oc ei ini as rise er eee C. rubidus
libPrecloacal sulcus ADSEMt: iirc oo. eases + aq sate coke eieueen © o.cicho) cleans ace olokclcr a ca ee 2)
2a. Greatly enlarged pore-bearing scale posterior to main precloacal pore row ...C. chrysopylos
ZbwNovenlarged:pore-bearing scales 5. 5 Hee ae aie oe aici le cee a nie ee een eee 3
3a. Ventral scales enlarged, equal in number to dorsal tubercle rows .............. C. aequalis
3b. Scale rows across mid-belly substantially exceed dorsal tubercle rows ................. 4+
4a. Males? with precloacal pores only (or precloacal pores barely reaching thigh base) ....... 5
4b. Males with diastema between precloacal and femoral pore series..............---+--- 14
Sapmemoraliscales enlarged, oy 5. < iene.e otis. g, «Famers oe 8 oat ate c genss Stone 6
SDmmemoraliscales not enlarged o.oo so ses coe aid Pmge 14> ere a Oe eee 10
GasDark dorsum patterned with light markings »..:.<.. 425.2255. 24) o> asec eee df
Gbwereht dorsum patterned with dark markings... .. 4.25: 2.52.2. .5>>9456 0 eee eee 8
7a. Light markings arranged transversely, head with light reticulations................ C. feae
7b. Light markings arranged longitudinally, head without reticulations ............ C. oldhami
Sa) Morsal pattern of thin dark bands with light edging’)....-.522-.>---+-5 5 se eee 9
SbyWorsalipattem! of dark’spots orregular blotches*> 2.22. . > ee sess lee eee C. peguensis
9) aac ~
~ Female condition only known for C. wakeorum
BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 495
SeeMOrsuMm owhcadimpattemedies eignce sire es Wy. Pepe em ee. C. wakeorum
SeePlorsum of head withidarkmarkines*7 052). onissee a.) . pis te eee oe C. consobrinoides
10a. Median subcaudal scales enlarged to form transverse plates ............... C. variegatus
MibBNORMe Gan SubCaudal plates icin a: seer cole eae ON pes aaa he Stay fa Lets Ce ete eT 11
PlaDicits very short, subcaudal scalation granular 2... Mes) sa5. 22 eae. C. brevidactylus
11b. Digits moderately long, subcaudal scales much larger than dorsal tail scales........... 12
12a. Precloacal pores in a strongly angled series, recessed in a shallow groove (males only);
FEMI eSaWilinpreclOacal: POLES PLESENU cj. ccs. seein suse Ns es ees scene C. gansi
12b. Precloacal pores less acutely angled, not recessed; pores absent in females............ 13
13a. Dorsal pattern of paired dark rectangular markings with white punctuations or bands, light
bandsxon:tailymuchimarrowen than darks 4834 ach: oe nl | eosin eke C. ayeyarwadyensis
13b. Dorsal pattern not as above, light and dark tail bands subequal. ............ C. khasiensis
Ierambearce(s Vilto over 1 00imm)sdicitsclongatess. 4) as- ee oee e ee 15
b= small(SVil< 56 mm); digits relativelysshort. Ji. (2 oe. ee ae. C. annandalei
15a. Precloacal pores 15, femoral pores 16—19 per thigh, dorsal pattern of transverse dark bands
30 5.550 Groh Ge antes CRD RCE RCE Ae ates Sars aA ERT Ree AE CEC eR SR rr ae C. russelli
15b. Precloacal pores 9-11, femoral pores 11—12 per thigh, dorsal pattern of regular, paired dark
| TILE CCSS wie ah a Ree eee Tne EN Aes Pe Fe geet Fett, WL Ra ERE C. slowinskii
To date a complete revision of Cyrtodactylus has not been attempted, nor have any phyloge-
netic analyses of the group been undertaken. Phenetic similarity, however, suggests some probable
affinities among the species of Myanmar. It is clear that C. russelli is closely allied to C. slowin-
skii. These are the only two species in Myanmar to exceed 100 mm SVL and the two share simi-
larities in precloacal and femoral pore configuration as well as color pattern. Together these species
are probably part of a more widespread clade of large-bodied, long-fingered forms that includes C.
consobrinus and other species extending from southeast Asia through the Indoaustralian
Archipelago to the western Pacific. Although less similar in overall appearance, C. aequalis and C.
chrysopylos also have elongate digits, relatively large body size and well-demarcated, denticulate
ventrolateral body folds and are probably members of this same clade.
Cyrtodactylus wakeorum is phenetically very similar to C. consobrinoides and is probably
closely related to this form. Cyrtodactylus annandalei is also somewhat similar in appearance to C.
consobrinoides, and to C. peguensis, but its pore configuration is substantially different from
either. All of these species are relatively small bodied and have moderately short digits and weak-
ly-developed ventrolateral folds, without denticulate margins. Cyrtodactylus ayeyarwadyensis and
C. gansi are very similar to C. khasiensis with respect to size, body proportion, scalation, and gen-
eral color pattern. These species almost certainly are each others closest relatives and replace one
another geographically from the northern Assam border ranges (C. khasiensis), to the Chin Hills
(C. gansi), to the lowlands west and south of the Rakhine Yoma (C. ayeyarwadyensis).
The distribution of the seven new species of Cyrtodactylus, and of all geckos in Myanmar, is
largely a function of collecting effort. That so many new taxa should be discovered in a short peri-
od reflects both the intensive collecting effort of the Myanmar Herpetological Survey and the fact
that the Survey has collected chiefly in areas that have not previously been explored herpetologi-
cally. For example, the collections made and reported on by Theobald (1868) were chiefly from
Pegu (Bago) and Tenasserim (Tanintharyi). Even Fea, who traveled extensively in both Upper and
496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 25
Lower Burma, including Tenasserin, Carin (now Kayin) State, Mandalay, and Bhamo (Banmo) in
the Cactin (Kakhien) Mountains near the Yunnan border (Boulenger 1887a, 1887b, 1888, 1893;
Fea 1897; Hallermann et al. 2002), collected almost exclusively east of the Ayeyarwady River.
It is not surprising that the only one of the species for which several localities over a distance
of more than 100 km exist is the lowland form C. ayeyarwadyensis. Although the additional local-
ities for the other taxa may be expected, it seems likely that most Cyrtodactylus in Myanmar will
be found to be restricted to individual montane areas or hill ranges. This is almost certainly the case
for C. brevidactylus, which occurs on the isolated Mt. Popa in north-central Myanmar, but proba-
bly also applies to species inhabiting the north-to-south running ranges both east and west of the
central dry zone, which is itself an apparent area of endemism (Bauer 2002). Despite the intensive
work of the Myanmar Herpetological Survey, Annandale’s (1905b) statement that “the country
between northern Assam and southern Tenasserim is one . .. which even the systematist has not yet
exhausted the vertebrate zoology” is as true today as it was nearly a century ago.
ACKNOWLEDGMENTS
Specimens examined in this paper were obtained by the Myanmar Herpetofaunal Survey, a
joint program of the Myanmar Nature and Wildlife Conservation Division, Forest Department, the
California Academy of Sciences, and the Smithsonian’s National Museum of Natural History with
support from the National Science Foundation (Grant DEB 9971861). I thank George R. Zug and
the late Joseph B. Slowinski for inviting me to work on the geckos collected by the Myanmar
Herpetofaunal Survey and all of the collectors noted herein for providing such a rich source of
material to work from. Robert C. Drewes, Alan E. Leviton and Jens V. Vindum (CAS), George Zug,
Robert Wilson and Ken Tighe (USNM) and James Hanken and José P. Rosado (MCZ) provided
access to specimens in their care. Michelle S. Koo kindly prepared the map and Guinevere O.U.
Wogan assisted in locating photographs of living geckos. Indraneil Das and Thomas Ulber assist-
ed with the literature.
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ANNANDALE, N. 1913. The Indian geckos of the genus Gymnodactylus. Records of the Indian Museum
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BAUER, A.M. 2002. Two new species of Cyrtodactylus (Squamata: Gekkonidae) from Myanmar. Proceedings
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BAUER: SEVEN NEW SPECIES OF CYRTODAYCYLUS FROM MYANMAR 497
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di Storia Naturale di Genova, ser. 2, 13:304-347, pls. 7-12.
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with a key to the species from Vietnam. Asiatic Herpetological Research 7:19—22.
Das, I. 1997. A new species of Cyrtodactylus from the Nicobar Islands, India. Journal of Herpetology
31:375-382.
Das, L, AND L.J. Lim. 2000. A new species of Cyrtodactylus (Sauria: Gekkonidae) from Pulau Tioman,
Malaysia. Raffles Bulletin of Zoology 48:223-231.
FEA, L. 1897. Viaggio di Leonardo Fea in Birmania e regioni vicine 76. Riassunto generale dei risultati zoo-
logici. Annali del Museo Civico di Storia Naturale di Genova, ser. 2, 17:385-658.
HALLERMANN, J., N. ANANJEVA, N. ORLOV AND F. TILLACK. 2002. Leonardo Fea’s historical collection of
Amphibia and Reptilia from Burma deposited at the Zoologisches Museum Hamburg. Mitteilungen aus
dem Hamburgischen Zoologischen Museum und Institut 99:139-153.
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tions of three new species. Japanese Journal of Herpetology 13:91—107.
Hora, S.L. 1926. Notes on lizards in the Indian Museum. I. On the unnamed collection of lizards of the
Family Geckonidae. Records of the Indian Musuem 28:187-193, pl. 7.
HUNDLEY, H.G. 1964 et seq. Check list of reptiles of Burma. Burmese Forestry Department internal document
[mimeograph], pp. 1-111, 1-28 [1964]; also Supplement I. Check list of reptiles of Burma, pp. 1-7 [ca.
1965].
INGER, R.F. 1999. Distribution of amphibians in southern Asia and adjacent islands. Pp. 445-482 in W.E.
Duellman, ed., Patterns of Distribution of Amphibians, a Global Perspective. Johns Hopkins University
Press, Baltimore.
KLuGE, A.G. 1983. Cladistic relationships among gekkonid lizards. Copeia 1983:465—475.
KiuGe, A.G. 1991. Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service
(85):1-35.
KLUGE, A.G. 1993. Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego.245 pp.
KiuGE, A.G. 2001. Gekkotan lizard taxonomy. Hamadryad 26:1—209.
ROsLER, H. 2000. Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia:
Gekkonomorpha). Gekkota 2:28-153.
Shreve, B. 1940. Reptiles and amphibians from Burma with descriptions of three new skincs. Proceedings of
the New England Zoological Club 18:17—26.
SLOWINSKI, J.B., AND W. WutsTer. 2000. A new cobra (Elapidae: Naja) from Myanmar (Burma).
Herpetologica 56:257-270.
SuiTH, M.A. 1935. The Fauna of British India,Including Ceylon and Burma. Reptilia and Amphibia. Vol. IL.-
Sauria. Vaylor and Francis, London. xiii + 400 + [2] pp., | pl., 2 folding maps.
SZCZERBAK, N.N., AND M.L. GOLUBEV. 1977. Systematics of the Palearctic geckos (genera Gymnodactylus,
Bunopus, Alsophylax). Proceedings of the Zoological Institute, Academy of Sciences of the USSR
74:120—-133. [in Russian] :
SZCZERBAK, N.N., AND M.L. GoLuBEV. 1984. On generic assignement of the Palearctic Cyrtodactylus lizard
species (Reptilia, Gekkonidae). Vestnik Zoologii 2:50—56. [in Russian]
SZCZERBAK, N.N., AND M.L. GOLUBEV. 1986. Gecko Fauna of the U.S.S.R. and Contiguous Regions. Naukova
Dumka, Kiev. 232 pp., 8 pp. pls. [in Russian]
THEOBALD, W. 1868. Catalogue of the reptiles of British Birma, embracing the provinces of Pegu, Martaban,
498 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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and Tenasserim; with descriptions of new or little-known species. The Journal of the Linnean Society
10:4-67.
ULBER, T. 1993. Bemerkungen tiber cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei
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London 124:469-492.
Copynght © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26, pp. 499-662, 97 figs., 33 maps, 2 tables November 14, 2003
Species Revision of the Coelotine Spider Genera
Bifidocoelotes, Coronilla, Draconarius, Femoracoelotes,
Leptocoelotes, Longicoelotes, Platocoelotes, Spiricoelotes,
Tegecoelotes, and Tonsilla (Araneae: Amaurobiidae)
Xin-Ping Wang!
Schlinger Foundation Postdoctoral Fellow and Research Associate
Department of Entomology, California Academy of Sciences
Golden Gate Park, San Francisco, CA 94118
Table of Contents
PAEIS CEA [PRN ST NE AT cA cI ste SI ETO OO ee Ek AIS ME AE a hey re 499
OURO ANTICO) VS sho ea ins ene Ne eae ee Ar Ae ee Re ba at 500
PEA CHidSTANGIWIECHOUS a tive he cre saras, field Pans or Lhe y CORAM MAME Ryne claret gy OMA, Men Avo ca 500
GI OWSG BATTS Raekh attains dealt geet nelle ey era bea Niet di hu aS gee tiaaR Gv Higste dt Me 501
DESOTO TIONG: aati ne ae See PO ret Rae ntact de i eit ea te ini cM enti tewiet s HhanN 502
AEP OCOCLOLESINV ATS, DOO 2. rr gens cet a) ore vate tale NCE Mesto ie Te ERE Lia area ee cae Gee 502
OKO TIA NN AIS OO A oregon anne kt pccten optckce Tan as EM CR Ra Li mass eet acter epa) eh ora ee 503
DF ACONAMUSION(C Hani KOV gh O99 OM en Macey dares eh Wy ears La eee eon ate 507
ICI ONACOCIOLESEWAN SOU 2 28 Pane t 1 py eee los) eee Eee 5p)
FEDLOCOCIOLESW AN Os ZOO 2 erat ih site nade het NONE oP cP oR AS poste eee LRM ee emo ae 558
WOU SICOCIOLES WALL Ong) ODM tok ir Rate Rec et te art state ponent a miscyin ae SERRA Sy)
TH ALOCOCIOLES NN ANS, LOO Dre eee ae se ead ue ey eae Se La Lenses hacen chey peerage 561
PLE COCLOLES INN AM Oot (NUD exer ie Pa heeh ode elec ee ea keg ey Sieh Sate oe rape ek ee 563
egecoclotesOvichinnikov, Tog Oe rink cee een ees ee annette ty i keane) Aer: 566
horisiilamanexand Nats WOO Dyck mcs crc genen et pak ees a eaten SON CEE apse Daren) a ns cries eke ep Neca aie 570
| ALSSALTUNE (CHSC, asta a ae ie aa eae te RG? gt alten Ane SSPE) Reig ars TaN nice AMORA Fac, 6 576
bssteatronssanGuMiapsrp a. tvcis Shee. ie Ske aoc ce) ese es Me UA I OS ia Se eset: 580
DELS fiat (POTS eens ey pecs Cuore Shee, ae ta ns ONDA wats Retr ANE E RNY URS TNR saan ea rau core 581
IVE APSE erent nas. « Can tenre ee Maat re ants IER: AT em cA caliet ca MULE tacks eater eee ae Ae 646
The following coelotine genera from East Asia are revised: Bifidocoelotes Wang, with
two species; Coronilla Wang, with five species, including two new species (C. libo and
C. subsigillata) and one new synonym (the species C. yanling Zhang and Yin, 2001 is
a junior synonym of C. gemata); Draconarius Ovtchinnikov, with 86 species, includ-
ing 24 new species (D. baxiantaiensis, D. haopingensis, D. episomos, D. absentis, D.
agrestis, D. capitulatus, D. curiosus, D. disgregus, D. dissitus, D. dubius, D. griswoldi,
D. incertus, D. linxiaensis, D. nudulus, D. parabrunneus, D. paraterebratus, D.
patellabifidus, D. pseudobrunneus, D. pseudocapitulatus, D. pseudowuermlii, D. rotun-
dus, D. simplicidens, D. tibetensis, and D. yichengensis), five new synonyms (the
species D. parawudangensis Zhang, Zhu and Song, 2002 is a junior synonym of D.
! Current Address: Illinois Natural History Survey, 167 E. Peabody Dr., Champaign, IL 61820
499
500 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
wudangensis, the species D. sinualis (Chen, Zhao and Wang, 1991) is a junior syn-
onym of D. lutulentus, the species Coelotes rufuloides Zhang, Peng and Kim, 1997 is
a junior synonym of D. rufulus, the species Coelotes shuangpaiensis Peng, Gong and
Kim, 1996 is a junior synonym of D. digitusiformis, and the species D. baccatus
(Wang, 1994) is a junior synonym of D. neixiangensis), and 37 new combinations, all
transferred from genus Coelotes (D. acidentatus (Peng and Yin, 1998), D. adligansus
(Peng and Yin, 1998), D. altissimus (Hu, 2001), D. amygdaliformis (Zhu and Wang,
1991), D. argenteus (Wang et al., 1990), D. bituberculatus (Wang et al., 1990), D. brun-
neus (Hu and Li, 1987), D. carinatus (Wang et al., 1990), D. chaigiaoensis (Zhang,
Peng and Kim, 1997), D. denisi (Schenkel, 1963), D. digitusiformis (Wang et al.,
1990), D. everesti (Hu, 2001), D. funiushanensis (Hu, Wang and Wang, 1991), D.
gyriniformis (Wang and Zhu, 1991), D. hangzhouensis (Chen, 1984), D. himalayaen-
sis (Hu, 2001), D. hui (Dankittipakul and Wang), D. huizhunesis (Wang and Xie,
1988), D. jiangyongensis (Peng, Gong and Kim, 1996), D. linzhiensis (Hu, 2001), D.
magniceps (Schenkel, 1936), D. nanyuensis (Peng and Yin, 1998), D. ornatus (Wang
et al., 1990), D. penicillatus (Wang et al., 1990), D. pervicax (Hu and Li, 1987), D. picta
(Hu, 2001), D. gingzangensis (Hu, 2001), D. quadratus (Wang et al., 1990), D. rufulus
(Wang et al., 1990), D. streptus (Zhu and Wang, 1994), D. strophadatus (Zhu and
Wang, 1991), D. subtitanus (Hu, 1992), D. syzygiatus (Zhu and Wang, 1994), D. tere-
bratus (Peng and Wang, 1997), D. tryblionatus (Wang and Zhu, 1991), D. uncinatus
(Wang et al., 1990), and D. yadongensis (Hu and Li, 1987); Femoracoelotes Wang,
with two species; Leptocoelotes Wang, with two species; Longicoelotes Wang, with
three species, including two new combinations, all transferred from genus Coelotes
(L. kulianganus (Chamberlin, 1924) and L. senkakuensis (Shimojana, 2000)); Plato-
coelotes Wang, with five species, including one new species (P. kailiensis); Spiri-
coelotes Wang, with three species, including one new species (S. pseudozonatus);
Tegecoelotes Ovtchinnikovy, with five species; and Tonsilla Wang and Yin, with seven
species, including one new species (7: makros) and two new combinations, all trans-
ferred from genus Coelotes (T. lyratus (Wang et al., 1990) and T. tautispinus (Wang et
al., 1990)).
Coelotine spiders are endemic to the Holarctic region, where they are distributed from eastern
North America, Europe, middle Asia, to East Asia, and comprise at least 277 species (Platnick
2000-2002; Wang 2002). Wang (2002) provided a generic revision and recognized 20 genera. The
ongoing species revision by the author shows a great coelotine species diversity, particularly in
East Asia.
The species of the North American genus Wadotes Chamberlin, 1925, with 11 described
species, were revised by Bennett (1987) and those of the Himalayan genus Himalcoelotes Wang,
2002, with 10 described species, were revised by Wang (2002). In the present study, 10 East Asian
genera are revised: they are Bifidocoelotes Wang, 2002; Coronilla Wang, 1994; Draconarius
Ovtchinnikoy, 1999; Femoracoelotes Wang, 2002; Leptocoelotes Wang, 2002; Longicoelotes
Wang, 2002; Platocoelotes Wang, 2002; Spiricoelotes Wang, 2002; Tegecoelotes Ovtchinnikov,
1999; and Tonsilla Wang and Yin, 1992.
MATERIALS AND METHODS
As coelotines are somatically relatively uniform, descriptions of the new species record only
variable structures, such as chelicerae, eyes, and male and female genitalic structures, with special
emphasis on genitalic structures in redescriptions of known species. Eyes, legs, and body lengths
are given in the new species descriptions.
WANG: EAST ASIA COELOTINE SPIDERS 501
All measurements are in mm. Eye sizes are measured as the maximum diameter from either
dorsal or frontal views. Leg measurements are given as: total length (femur, patella + tibia, metatar-
sus, tarsus). All scale lines are 0.2 mm length except where indicated otherwise.
ABBREVIATIONS.— ALE — anterior lateral eyes; AME — anterior median eyes; PLE — posteri-
or lateral eyes; PME — posterior median eyes; RTA — retrolateral tibial apophysis.
ACKNOWLEDGMENTS
I thank Charles Griswold (CAS) and Norman Platnick (AMNH) for their continuing support.
Darrell Ubick and Charles Griswold of CAS critically read the manuscript in draft. A Schlinger
Foundation (CAS) supported my study as a postdoctoral research fellow in the Department of
Entomology at the California Academy of Sciences. Additional support for this research came from
the China Natural History Project of the California Academy of Sciences (CAS) and the US
National Science Foundation grant DEB 0103795. This is Scientific Contribution no. 26 from the
California Academy of Sciences Center for Biodiversity Research and Information (CBRI) and
contribution no. 19 from the China Natural History Project (CNHP).
I am especially grateful to the following curators and institutions that loaned East Asian mate-
rial for this study:
AMNH — American Museum of Natural History, New York, USA. N.I. Platnick
AMNH-CU — Cornell University Collection loaned to the AMNH. N.I. Platnick
BMNH -— The Natural History Museum, London, UK. P. Hillyard and J. Margerison
CAS — California Academy of Sciences, California, USA. C.E. Griswold
CSO — Collection of Ovtchinnikov, Bishkek, Kyrghyzstan. $.V. Ovtchinnikov
HBI — Hunan Biological Institute, Changsha, China. X.J. Peng and C.M. Yin
HEC — Hope Entomological Collections, Oxford, England. M. Akinson
HTC — Hangzhou Teachers College, Hangzhou, China. Z.F. Chen
HTU — Hebei Teachers University, Shijiazhuang, China. M.S. Zhu
HUB — Hebei University, Baoding, China. M.S. Zhu
HUW — Hubei University, Wuhai, China. J. Chen and J.Z. Zhao
IZB — Institute of Zoology, Beijing, China. J. Chen and D.X. Song
KAI — Korean Arachnological Institute, Seoul, Korea. J.P. Kim
MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA.
L. Leibensperger.
MNHN — Museum National d’ Histoire Naturelle, Paris, France. C. Rollard
NBUMS — Norman Bethune University of Medical Sciences, Changchun, China. J.C. Gao and C.D. Zhu
NHMB -— Naturhistorisches Museum Basel, Basel, — Switzerland. A. Hinggi
NRS — Naturhistoriska Riksmuseum, Stockholm, Sweden. T. Kronestedt
NSMT — National Science Museum, Tokyo, Japan. H. Ono
PSU — Perm State University, Russia. V. Efimik and S. Esyunin
SMF — Senckenberg Museum, Frankfurt, Germany. M. Grasshoff and J. Martens
SZM — Siberian Zoological Museum, Novosibirsk, Russia. D.V. Logunov
THU — Department of Biology, Tunghai University, Taichung, Taiwan. I.M. Tso
USNM — National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA.
J. Coddington
ZMB — Museum fur Naturkunde, Zentralinstitut der Humboldt-Universitut zu Berlin, Berlin, Germany.
J. Dunlop and Sh. Nawai.
ZSM — Zoologische Staatssammlung, Miinchen, Germany. E. Karl
IZ] — Institute of Zoology, Innsbruck, Austria. K. Thaler
Institution abbreviations used in this paper also include:
SDU — Shandong University, Jinan, China.
502 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
‘TAXONOMY
Genus Bifidocoelotes Wang, 2002
Bifidocoelotes Wang, 2002:37 (type species, by original designation, Coelotes bifida Wang, Tso and Wu,
2001, from Taiwan).
DIAGNOsIS.— The female can be distinguished from other coelotines by having the single,
bifurcate epigynal tooth. The male is similar to Asiacoelotes and Draconarius by having the elon-
gated cymbial furrow and long embolus, but differs from Asiacoelotes by the presence of a con-
ductor dorsal apophysis and from Draconarius by the broad, bifurcate conductor and the small
median apophysis (Figs. 1—2).
PHYLOGENETIC PLACEMENT.— Remains unresolved; with genera Draconarius, Asiacoelotes,
and the Platocoelotes + Spiricoelotes clade, supported by the strongly elongated spermathecal
tubes, well-developed cymbial furrow, long embolus, and posteriorly originating embolic base
(Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China (Map 1).
COMPOSITION.— 2 species:
1. Bifidocoelotes bifidus (Wang, Tso and Wu, 2001)
2. Bifidocoelotes primus (Fox, 1937)
Bifidocoelotes bifidus (Wang, Tso and Wu, 2001)
Figures 1A—D; Map 1
Coelotes bifida Wang, Tso and Wu, 2001:128, figs. 1-10 (female holotype and male paratype from Nantou,
Taiwan, in THU, examined).
Bifidocoelotes bifida: Wang, 2002:38, figs. 86—100.
DIAGNosIs.— Distinguished from B. primus by the shorter epigynal bifurcation (less then 2
of the total length), the strongly developed copulatory ducts, the medially situated spermathecal
heads, and the widely separated, longitudinally elongated spermathecae (Figs. 1A—D).
DESCRIPTION.— Described by Wang, Tso and Wu (2001) and Wang (2002). Chelicerae with
three promarginal and two retromarginal teeth. Female epigynum with single, long, slightly bifur-
cate epigynal tooth (less than 2 total length); atrium small, anteriorly situated, near base of epigy-
nal tooth; copulatory ducts long, strongly convoluted mesad of spermathecae, and slightly extend-
ing laterad of spermathecae; spermathecal heads small, situated anteriorly, close together; sper-
mathecal bases widely separated, stalks elongated, slightly convoluted, widely separated and ante-
riorly converging (Figs. |A—B). Male palp with patellar apophysis large, with slightly curved apex;
RTA long; lateral tibial apophysis small; cymbial furrow longer than half cymbial length; conduc-
tor broad, more or less spiraled, with bifurcate apex; conductor with dorsal edge bearing broad
membranous extension, dorsal apophysis small, lamella well developed; embolus posterior in ori-
gin, long, slender; median apophysis small, spoon-like (Figs. 1C—D).
DISTRIBUTION.— China (Taiwan) (Map. 1).
MATERIAL EXAMINED.— CHINA: Taiwan: Nantou County, Huei-Sun Experimental Forest, ele-
vation 1680m, March 31, 1998, female holotype (Hai-Yin Wu; THU, THU-Ar-990017); Nantou
County, Huei-Sun Experimental Forest, March 31, 1998, 1 male paratype (Hai-Yin Wu; THU,
THU-Ar-990020); Nantou County, Huei-Sun Experimental Forest, elevation 1680m, March 31,
1998, 1 male (Hai-Yin Wu, THU, THU-Ar-990019); Nantou County, Huei-Sun Experimental
Forest, elevation 1675m, March 31, 1998, 1 male (Hai-Yin Wu, THU, THU-Ar-990026): Nantou
WANG: EAST ASIA COELOTINE SPIDERS 503
County, Huei-Sun Experimental Forest, elevation 1550m, March 31, 1998, 1 female (Hai-Yin Wu,
THU, THU-Ar-990027).
Bifidocoelotes primus (Fox, 1937)
Figures 2A—B; 97G; Map 1
Wadotes primus Fox, 1937:1, figs. 1-2 (female holotype and female paratype from Hong Kong, China, in
AMNH and USNM, examined). Bennett, 1987:126, figs. 109-110.— Song, Zhu and Chen, 1999:395.
Bifidocoelotes primus: Wang, 2002:37.
DIAGNosis.— Distinguished from B. bifidus by the strongly bifurcate epigynal tooth (bifurca-
tion more then 2 total length), the short copulatory ducts, the laterally situated, medially extend-
ing spermathecal heads, the closely situated spermathecae, and the transversely extending sper-
mathecal bases (Figs. 2A—B).
DESCRIPTION.— Described by Fox (1937). Chelicerae with three promarginal and two retro-
marginal teeth. Female epigynum with single bifurcate tooth, bifurcation more than one half total
length; atrium broad; epigynal hoods deep, situated laterad of atrium; copulatory ducts short; sper-
mathecal heads long, slender, originating laterad of spermathecae, anteriorly extending and con-
verging; spermathecal bases large, close together, laterally extending, stalks broad, convoluted, sit-
uated close together (Figs. 2A—B).
DISTRIBUTION.— China (Hong Kong) (Map. 1)
MATERIAL EXAMINED.— CHINA: Hong Kong: Tingping Mt., October 14, 1997, 1 female (X.
P. Wang, IZB); Hong Kong, female holoytpe (AMNH); Hong Kong, late January, 1920, | female
paratype (Bassett Digby, USNM).
Genus Coronilla Wang, 1994
Coronilla Wang, 1994: 281 (type species, by original designation, Coronilla gemata Wang, 1994, from
China).— Platnick, 1997:667;— Wang, 2002:61.
DIAGNOSIS.— The male can be distinguished from all other coelotine genera except
Femoracoelotes by the absence of a lateral tibial apophysis, and from Femoracoelotes by the
absence of a femoral apophysis and the presence of two patellar apophyses. The female can be rec-
ognized by the broad atrium, the presence of transversely extending atrial carina, and the absence
of epigynal teeth (Figs. 3-6).
PHYLOGENETIC PLACEMENT.— The presence of four cheliceral retromargin teeth suggests
Coronilla is the sister group of Femoracoelotes from Taiwan; together they form the sister group
of all other coelotines (Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China, Vietnam (Map 2).
COMPOSITION.— 5 species, including 2 new species and | new synonym:
1. Coronilla gemata Wang, 1994
Coelotes huangsangensis Peng et al., 1998
Coelotes yoshikoae Nishikawa, 1995
Coronilla yanling Zhang and Yin, 2001, NEw SYNONYMY
. Coronilla libo, sp. nov.
. Coronilla mangshan Zhang and Yin, 2001
. Coronilla sigillata Wang, 1994
. Coronilla subsigillata, sp. nov.
A BW bh
504 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Key To the Species of the Genus Coronilla
leeNialex(those of C. subsigillatajunknown). 55 .024..- 2) aes) eee D,
Remale (those of Gylibo unknown)... 56:04. eca% © cia meee: < ae obec el or eee 5
PmaatellanwathnthnneerapOPMYSES:ccc es. cc). 56s awe gg ais os ee ae ee mangshan
Patellaiwith tworapophyses. 2... 2% 5... ole aoa goes Soe ele a eee 3)
3. Median apophysis spiraled; conductor simple, not bifurcate, without ventral apophysis (Fig. 4B)
4 A nee are <td pene Eek eee aurea MEAP AMO 5 oo oo 0.000 20 6 libo
Median apophysis not spiraled; conductor either bifurcate (Fig. 5D) or with ventral apophysis
(EIGASD)E eececee cota wets Sancuso ote si ohs etry GO Sie oe epee a, oO 4
4. Ventral patellar apophysis slender; conductor with large ventral apophysis; conductor dorsal
apophysis'small;not toothed (Figs: 3€-E)5. 2... 28 a5 552 ee ee gemata
Ventral patellar apophyses broad; conductor with small ventral apophysis; Conductor dorsal
apophysisilarzetoothedi(Figs.5C=D) 22s 56 ae. oe. Sole eee ee eee sigillata
5. Spermathecae with laterally extending apophyses; spermathecal heads short, broad (Fig. 3B).
eS es ee AEST Sea aE Re aE eC PERM UNM MBS SSS Goo a oo gemata
Spermathecae without laterally extending apophyses; spermathecal heads long, slender (Figs.
SB SOB) hice. sae leasioes be ales Sars toe ge neaeaeene Gece dl Sie? gh gins Sty > Cee eee 6
Om Copulatory ductsoundeds anteriorly situated) ae ee ee mangshan
Copulatory ducts posteriorly extending... 5 7 5. et ns oe ee ee 7
7. Copulatory ducts with broad, slightly lobed posterior edges; spermathecal heads originating
dorsalllyx(Eig SB) Neen 5 fst cct a Boor iie ccs, aeaseaes creek nevete cP cael «oe ane eC eae sigillata
Copulatory ducts with narrow, non-lobed posterior edges; spermathecal heads originating ven-
tralllyi(BignGB) nn er reat eee S46 Goleta d Basses Oe Seoece veers Shahi e eee subsigillata
Coronilla gemata Wang, 1994
Figures 3A—E; Map 2
Coronilla gemata Wang, 1994:281, figs. 1-5 (female holotype and male allotype from Mt. Zhangjiajie,
Dayong, Hunan, China, in HTU, examined).— Song, Zhu and Chen, 1999:389, figs. 229G—H, K-M;—
Wang, 2002:61, figs. 158-180.
Coelotes yoshikoae Nishikawa, 1995:141, figs. 1-8 (types from Vinh Phu Provy., Vietnam, in NSMT, not
examined).
Coelotes huangsangensis Peng et al., 1998:77, figs. 1-6 (female holotype and male allotype from Huangsang,
Suining, Hunan, China, in HBI, examined).
Coronilla yanling Zhang and Yin, 2001:489, figs. 8-11 (male holotype and | male paratype from Taoyuan-
dong, Yanling, Hunan, China, in HBI, not examined). NEw SYNONYMY.
DIAGNOsIs.— The male can be distinguished from C. libo by the presence of broad conductor
ventral apophysis and from C. sigillata by the slender patellar ventral apophysis. The female can
be recognized by the presence of spermathecal lateral apophyses and the anteriorly situated sper-
mathecal heads (Figs. 3A—E).
DESCRIPTION.— Described by Wang (1994) and Wang (2002). Chelicerae with three promar-
ginal and four retromarginal teeth. Female epigynum without teeth; atrium large, carina broad,
transversely extending: copulatory ducts large; spermathecal heads short, situated anteriorly; sper-
WANG: EAST ASIA COELOTINE SPIDERS 505
mathecal bases transversely extending, situated close together; spermathecal stalks broad, with lat-
eral apophyses (Figs. 3A—B). The male palp with two patellar apophyses, ventral slender and long
(occasionally short), dorsal short and strong; RTA slightly shorter than tibia; lateral tibial apoph-
ysis absent; cymbial furrow short; conductor short, ventral apophysis long, broad, anteriorly
curved; conductor dorsal apophysis large; median apophysis slender, with slightly curved apex
(Figs. 3C—E).
DISTRIBUTION.— China (Hunan, Sichuan) and VIETNAM (Vinh Phu) (Map 2).
MATERIAL EXAMINED.— CHINA: Hunan: Dayong, Mt. Zhangjiajie, November 5, 1985, female
holotype, 6 female and 8 male paratypes (J.F. Wang, HTU); Tianpingshan, October 16, 1986, 4
females and 4 males (J.R. Wang, HTU); Suining, Huangsang, October 14, 1996, female holotype
and male paratype of C. huangsangensis Peng et al., 1998 (M.X. Liu, HBI). Sichuan: E-mei-shan,
September 27, 1975, 2 females (C.D. Zhu, NBUMS, 75-2172); Chongqing, September 26, 1997,
molted to adult later October, 1 male (X.P. Wang, AMNH).
Coronilla libo Wang, sp. nov.
Figures 4A—B: Map 2
Typrs.— The male holotype, 2 male paratypes from Libo, Guizhou, China (October 5, 1997;
X. P. Wang), deposited in AMNH (holotype) and IZB (paratypes).
ETYMOLOGyY.— The specific name refers to the type locality.
DIAGNOsIS.— The male can be distinguished from C. gemata and C. sigillata by the simple
conductor (not bifurcate, without ventral apophysis), and the strongly spiraled median apophysis
(Figs. 4A—C).
MALE.— Total length 8.20. Carapace 4.20 long, 2.96 wide. Cheliceral promargin with three
teeth, retromargin four (occasionally five). Eye sizes and interdistances: AME 0.20, ALE 0.24,
PME 0.20, PLE 0.21, AME-AME 0.10, AME-ALE 0.05, PME-PME 0.13, PME-PLE 0.19, ALE-
PLE 0.04, AME-PME 0.18. Clypeal height 0.30. Leg measurements: I: 14.7 ( 4.00, 4.96, 3.84,
1.88); II: 13.1 (3.60, 4.41, 3.40, 1.64); HI: 11.6 (3.28, 3.80, 3.00, 1.48); IV: 14.6 (4.12, 4.72, 4.04,
1.68). Male palp with two patellar apophyses, ventral one relatively long and dorsally curved, dor-
sal one short; RTA slightly shorter than tibia; conductor slender, without ventral apophysis; con-
ductor dorsal apophysis broad; median apophysis slender, long, with spiraled apex (Figs. 4A—C).
FEMALE.— Unknown.
DISTRIBUTION.— China (Guizhou) (Map 2).
OTHER MATERIAL EXAMINED.— None.
Coronilla mangshan Zhang and Yin, 2001
Map 2
Coronilla mangshan Zhang and Yin, 2001:487, figs. 1-7 (male holotype, 1 male and 2 female paratypes from
Mangsha, Yizhang, Hunan, China, in HBI, not examined).
DIAGNOSIS.— The male can be distinguished from other species by the presence of three patel-
lar apophyses and the female by the rounded, anteriorly situated copulatory ducts.
DESCRIPTION.— Described by Zhang and Yin (2001). Cheliceral promargin with three teeth,
retromargin four. Female epigynum lacking teeth; atrium large; posterior margin strongly extend-
ing; copulatory ducts broad, rounded, anteriorly situated; spermathecal heads small, situated medi-
ally on spermathecae, close together; spermathecal bases small, slightly separated; spermathecal
506 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
stalks close together, anteriorly diverging. Male palp with three patellar apophyses; conductor with
ventral apophysis short, dorsal apophysis broad; median apophysis slender, slightly spiraled.
DISTRIBUTION.— China (Hunan) (Map 2).
MATERIAL EXAMINED.— None.
Coronilla sigillata Wang, 1994
Figures 5; Map 2
Coronilla sigillata Wang, 1994: 282, figs. 6-10 (female holotype, 2 male paratypes from Mt. Tianmushan,
Zhejiang, China, in HTU, examined).— Song, Zhu and Chen, 1999:389, figs. 229I-J, L-M.
NorTes.— Two male (one without palp) and three female types are examined from the same
vial labeled as C. sigillata, but only one female matches the illustrations of the holotype female
(Wang, 1994: figs. 9, 10). The two other females have quite different genitalic morphology and are
treated in this study as a new species, C. subsigillata sp. nov. All specimens were collected from
October 21 to October 23, 1974. It is possible that two Coronilla species exist in this location (Mt.
Tianmushan). Whether the males are C. sigillata or C. subsigillata sp. nov. is uncertain. Here in
this paper, the male specimens examined provisionally associated with C. sigillata.
DIAGNOsISs.— The male can be distinguished from other species by one strongly developed
and one much reduced patellar apophysis and the broad, toothed conductor dorsal apophysis (Figs.
5C-E). The female can be recognized from C. subsigillata by the posteriorly notched copulatory
ducts and the differences between their spermathecal head shapes (Figs. 5A—B).
DESCRIPTION.— Described by Wang (1994). Cheliceral promargin with three teeth, retromar-
gin four. Epigynal teeth absent; atrium large, carina broad, transversely extending; posterior mar-
gin strongly extending; copulatory ducts broad, with notched posterior end; spermathecal heads
long, slender, situated dorsally on spermathecae; spermathecal bases small, situated close togeth-
er; spermathecal stalks long, wide apart anteriorly and converging posteriorly (Figs. 5A—B). Male
palp with two patellar apophyses, ventral long and strong, dorsal short; RTA as long as tibial length;
conductor with ventral apophysis long, dorsal apophysis broad, with sharp anterior tooth and
rounded posterior process; median apophysis relatively broad, with slightly curved apex (Figs.
5C-E).
DISTRIBUTION.— China (Zhejiang) (Map 2).
MATERIAL EXAMINED.— CHINA: Zhejiang: Mt. Tiamushan, October 21—23, 1974, female holo-
type and 2 male paratypes (J.R. Wang, HTU).
Coronilla subsigillata Wang, sp. nov.
Figures 6, Map 2
TyPES.— Female holotype from Tianmushan, Zhejiang, China (October 21-23, 1974; J.F.
Wang), deposited in HTU.
ETYMOLOGY.— The specific name refers to its similarity to C. sigillata.
DIAGNOSIS.— This new species is similar to C. sigillata but can be distinguished by the less
expanded posterior atrial margin, the posteriorly extending, non-notched copulatory ducts, and the
ventrally situated spermathecal heads of the female (Figs. 6A—B).
FEMALES.— Total length 12.6. Carapace 5.80 long, 4.00 wide. Cheliceral promargin with three
teeth, retromargin four. Eye sizes and interdistances: AME 0.24, ALE 0.27, PME 0.22, PLE 0.23,
AME-AME 0.15, AME-ALE 0.15, PME-PME 0.29, PME-PLE 0.40, ALE-PLE 0.12, AME-PME
WANG: EAST ASIA COELOTINE SPIDERS 507
0.29. Clypeal height 0.42. Leg measurements: I: 15.3 ( 4.36, 5.36, 3.60, 2.00); HI: 13.5 ( 3.92, 4.64,
3.32, 1.64); I: 12.1 ( 3.48, 3.96, 3.12, 1.52); IV: 14.9 ( 4.24, 5.00, 3.96, 1.72). Epigynal teeth
absent; atrium large, carina broad, transversely extending; epigynum with posterior margin slight-
ly extending posteriorly; copulatory ducts broad, with extending, unnotched posterior ends; sper-
mathecal heads ventrally situated, mostly covered by copulatory ducts from dorsal view; spermath-
ecal bases small, situated close together; spermathecal stalks long, laterally extending (Figs.
6A-B).
MALE.— Unknown.
DISTRIBUTION.— China (Zhejiang) (Map 2)
OTHER MATERIAL EXAMINED.— CHINA: Zhejiang: Mt. Tiamushan, October 21—23, 1974, 1
female (not in good condition) (J.-F. Wang, HTU).
Genus Draconarius Ovtchinnikov, 1999
Draconarius Ovtchinnikov, 1999:70 (type species, by original designation, Draconarius venustus Ovtchinni-
kov, 1999 from Tadzhikistan).— Wang, 2002:66.
DIAGNOsIs.— Similar to Asiacoelotes in having an elongated cymbial furrow (longer than half
cymbial length) but can be distinguished by the posteriorly originating copulatory ducts and the
presence of a conductor dorsal apophysis; similar to Coelotes in having a patellar apophysis and a
conductor dorsal apophysis but can be distinguished by the elongated cymbial furrow (more than
half cymbial length), the long, posteriorly extending embolus, and the elongated median apophysis
of male and by the large copulatory ducts and the long spermathecae of female (Figs. 7-69).
PHYLOGENETIC PLACEMENT.— Remains unresolved with Bifidocoelotes, Asiacoelotes, and
the Platocoelotes+Spiricoelotes clade. Together they are supported by the strongly elongated sper-
mathecal tubes, the well-developed cymbial furrow, the long embolus, and the posteriorly originat-
ing embolic base (Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— Tadzhikistan, Bhutan, Nepal, China, Korea (Map 33).
GENITALIC VARIATIONS.— Patellar apophysis present in general, but can be absent in some
species; RTA long, occupying most of tibial length, or occasionally short, about half tibial length;
lateral tibial apophysis present; cymbial furrow long, broad, occupying more than half cymbial
length in most species, but can be short, less than half cymbial length; conductor simple, with dor-
sal apophysis; conductor lamella large in general, or reduced in some species; median apophysis
spoon-like, strongly elongated as in most species, but may be simple, not spoon-like, or absent;
embolus long, posterior in origin in most species, or short, prolateral in origin in others. Epigynal
teeth short in almost all species, but long as in huizhunesis group; epigynal teeth widely separated
in most species, but may be closely and anteriorly situated, or absent in few others; atrium small,
situated posteriorly in most species, but may vary in its shape and position in some species; copu-
latory ducts originated posteriorly, extending mesad of spermathecae in general, but may be
extending laterad of spermathecae, extending anteriorly, looping around spermathecae, or stongly
looped laterad of spermathecae; spermathecae broad, long in general, but can be short, rounded as
in gurkha group species.
SPECIES GROUPS.— Many Draconarius species are described in recent years with only male
or female. Most of those described with both male and female are only based on a limited number
of individuals and some might be incorrectly matched. As a result, a phylogenetic analysis at this
moment can hardly be achieved and in this study 35 of the 86 species are grouped only arbitrarily
based on the female genitalic characteristics; the other 51 are unplaced.
508 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
COMPOSITION.— 86 species, including 24 new species, 5 new synonyms, and 37 new combi-
nations. Among them, 35 species are grouped into 7 species groups according to female genitalia,
and the other 51 species remain unplaced:
The venustus group species
The female epigynum with epigynal teeth short, may be anteriorly and closely situated (Fig.
17A), or medially situated and widely separated (Figs. 13A, 67A), or accasionally lacking (Fig.
62A); copulatory ducts short, situated mesad of spermathecae; spermathecae broad, with short,
broad anterior expansion; spermathecal heads medially situated (Figs. 13B, 67B). Chelicerae with
three promarginal and two retromarginal teeth. Widespread in East Asia.
1. Draconarius aspinatus (Wang et al., 1990)
2. Draconarius baxiantaiensis, sp. NOV.
3. Draconarius calcariformis (Wang, 1994)
4. Draconarius colubrinus Zhang, Zhu and Song, 2002
5. Draconarius coreanus (Paik and Yaginuma, 1969) (In Paik, Yaginuma, and Namkung, 1969)
6. Draconarius davidi (Schenkel, 1963)
7. Draconarius funiushanensis (Hu, Wang and Wang, 1991), NEw COMBINATION (from Coelotes)
8. Draconarius gyriniformis (Wang and Zhu, 1991), NEw COMBINATION (from Coelotes)
9. Draconarius hui (Dankittipakul and Wang, 2003), NEW COMBINATION (from Coelotes).
Replacement name for preoccupied. Coelotes wangi Hu, 2001
10. Draconarius linzhiensis (Hu, 2001), NEw COMBINATION (from Coelotes)
11. Draconarius picta (Hu, 2001), NEw COMBINATION (from Coelotes)
12. Draconarius gingzangensis (Hu, 2001), NEW COMBINATION (from Coelotes)
13. Draconarius stemmleri (Brignoli, 1978)
14. Draconarius striolatus (Wang et al., 1990)
15. Draconarius trifasciatus (Wang and Zhu, 1991)
16. Draconarius venustus Ovtchinnikoy, 1999
17. Draconarius wudangensis (Chen and Zhao, 1997)
Draconarius parawudangensis Zhang, Zhu and Song, 2002, NEw SYNONYMY
18. Draconarius yadongensis (Hu and Li, 1987), NEW COMBINATION (from Coelotes)
19. Draconarius yostianus (Nishikawa, 1999)
The labiatus group species
The female epigynum with epigynal teeth short, anteriorly and closely situated (Figs. 37A,
66A); copulatory ducts short, situated mesad of spermathecae; spermathecae broad, with long,
strongly convoluted anterior expansion; spermathecal heads medially situated (Figs. 37B, 66B).
Chelicerae with three promarginal and two retromarginal teeth. Recorded from southern China.
20. Draconarius labiatus (Wang and Ono, 1998)
21. Draconarius wenzhouensis (Chen, 1984)
The lutulentus group species
The female epigynum with epigynal teeth short, widely separated (Figs. 39A, 40A) or absent
(Figs. 33A, 44A); copulatory ducts long, looping around spermathecae; spermathecae strongly
elongated and convoluted, anteriorly converging; spermathecal heads situated distally (Figs. 33B,
39B, 40B, 44B). Chelicerae with three promarginal and three retromarginal teeth. Distributed in
China and Himalayan region.
22. Draconarius haopingensis, sp. nov.
23. Draconarius lutulentus (Wang et al., 1990)
Draconarius sinualis (Chen, Zhao and Wang, 1991), NEw SYNONYMY
24. Draconarius molluscus (Wang et al., 1990)
25. Draconarius wuermlii (Brignoli, 1978)
WANG: EAST ASIA COELOTINE SPIDERS 509
The gurkha group species
The female epigynum with epigynal teeth short, widely separated (Figs. 30A, 32A); copulato-
ry ducts short; spermathecae short, rounded; spermathecal heads situated anteriorly (Figs. 30B,
32B: Hu 2000, figs. 845.2, 8—57.2). Chelicerae with three promarginal and two retromarginal
teeth. Recorded from Himalayan region.
26. Draconarius altissimus (Hu, 2001), NEw COMBINATION (from Coelotes)
27. Draconarius episomos, sp. Nov.
28. Draconarius gurkha (Brignoh, 1976)
Coelotes lama Brignoli, 1976
29. Draconarius himalayaensis (Hu, 2001), NEW COMBINATION (from Coelotes)
30. Draconarius subtitanus (Hu, 1992), NEw COMBINATION (from Coelotes)
The huizhunesis group species
The female epigynum with epigynal teeth strongly elongated, with slightly separated bases and
diverging apexes (Wang and Xu 1988, fig. 1; Zhu and Wang 1991, fig. 12); spermathecae long,
strongly convoluted (Wang and Xu 1988, fig. 2; Zhu and Wang 1991, fig. 13). Chelicerae with
three promarginal and two retromarginal teeth. Collected from eastern China.
31. Draconarius huizhunesis (Wang and Xie, 1988), NEw COMBINATION (from Coelotes)
C. huizhuneesis: Wang and Xie, 1988
C. huizhouensis: Song, Zhu and Chen, 1999
32. Draconarius strophadatus (Zhu and Wang, 1991), NEW COMBINATION (from Coelotes)
The terebratus group species
The female epigynum lacking epigynal teeth (Fig. 63A); copulatory ducts broad, anteriorly
extended; spermathecae short; spermathecal heads strongly elongated (Fig. 63B). Chelicerae with
three promarginal and two retromarginal teeth. Distributed in eastern and southern China.
33. Draconarius ornatus (Wang et al., 1990), NEw COMBINATION (from Coelotes)
34. Draconarius terebratus (Peng and Wang, 1997), NEw COMBINATION (from Coelotes)
The rufulus group species
The female epigynum lacking epigynal teeth; atrium small, posteriorly situated (Fig. 57A);
copulatory ducts broad, strongly convoluted with four to five loops laterad of spermathecae; sper-
mathecae long; spermathecal heads small (Fig. 57B). Chelicerae with three promarginal and two
retromarginal teeth. Distributed in eastern China.
35. Draconarius rufulus (Wang et al., 1990), NEw COMBINATION (from Coelotes)
Coelotes rufuloides Zhang, Peng and Kim, 1997, NEw SYNONYMY
Other non-grouped species:
36. Draconarius absentis, sp. nov.
37. Draconarius acidentatus (Peng and Yin, 1998), NEW COMBINATION (from Coelotes)
38. Draconarius adligansus (Peng and Yin, 1998), NEw COMBINATION (from Coelotes)
39. Draconarius agrestis, sp. Nov.
40. Draconarius amygdaliformis (Zhu and Wang, 1991), NEw COMBINATION (from Coelotes)
41. Draconarius arcuatus (Chen, 1984)
42. Draconarius argenteus (Wang et al., 1990), NEw COMBINATION (from Coelotes)
43. Draconarius baronii (Brignoli, 1978)
44. Draconarius bituberculatus (Wang et al., 1990), NEw COMBINATION (from Coelotes) -
45. Draconarius brunneus (Hu and Li, 1987), NEw COMBINATION (from Coelotes)
46. Draconarius capitulatus, sp. nov.
47. Draconarius carinatus (Wang et al., 1990), NEw CoMBINATION (from Coelotes)
48. Draconarius chaigiaoensis (Zhang, Peng and Kim, 1997), NEw CoMBINATION (from Coelotes)
49. Draconarius cheni (Platnick, 1989)
510
NO
50.
Sul
Sp
SS)
54.
SDs
0)
Ss
58.
ao
60.
61.
62.
63%
64.
65.
66.
67.
68.
69.
10:
qiile
WE
73:
74.
1B
76.
Ws
78.
WS
80.
81.
82
Epig
Epig
Epig
Epig
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Coelotes saxatilis Chen, 1984
Draconarius curiosus, Sp. NOV.
Draconarius denisi (Schenkel, 1963), NEW COMBINATION (from Coelotes)
Draconarius digitusiformis (Wang et al., 1990), NEw COMBINATION (from Coelotes)
Coelotes shuangpaiensis Peng, Gong and Kim, 1996, NEw SYNONYMY
Draconarius disgregus, sp. NOV.
Draconarius dissitus, sp. NOV.
Draconarius dubius, sp. nov.
Draconarius everesti (Hu, 2001), NEw COMBINATION (from Coelotes)
Draconarius griswoldi, sp. nov.
Draconarius hangzhouensis (Chen, 1984), NEW COMBINATION (from Coelotes)
Draconarius incertus, sp. Nov.
Draconarius infulatus (Wang et al., 1990)
Draconarius jiangyongensis (Peng, Gong and Kim, 1996), NEw CoMBINATION (from Coelotes)
Draconarius linxiaensis, sp. NOV.
Draconarius magniceps (Schenkel, 1936), NEW COMBINATION (from Coelotes)
Draconarius nanyuensis (Peng and Yin, 1998), NEW COMBINATION (from Coelotes)
Draconarius neixiangensis (Hu, Wang and Wang, 1991)
Draconarius baccatus (Wang, 1994), NEW SYNONYMY
Draconarius nudulus, sp. nov.
Draconarius parabrunneus, sp. nov.
Draconarius paraterebratus, sp. nov.
Draconarius patellabifidus, sp. nov.
Draconarius penicillatus (Wang et al., 1990), NEW COMBINATION (from Coelotes)
Coelotes penicilatus: Song, Zhu and Chen, 1999
Draconarius pervicax (Hu and Li, 1987), NEw COMBINATION (from Coelotes)
Draconarius potanini (Schenkel, 1963)
Draconarius pseudobrunneus, sp. nov.
Draconarius pseudocapitulatus, sp. Nov.
Draconarius pseudowuermilii, sp. nov.
Draconarius quadratus (Wang et al., 1990), NEw COMBINATION (from Coelotes)
Draconarius rotundus, sp. nov.
Draconarius schenkeli (Brignoli, 1978)
Draconarius simplicidens, sp. nov.
Draconarius singulatus (Wang et al., 1990)
Draconarius streptus (Zhu and Wang, 1994), NEw COMBINATION (from Coelotes)
. Draconarius syzygiatus (Zhu and Wang, 1994), NEw COMBINATION (from Coelotes)
83.
84.
85.
86.
Draconarius tibetensis, sp. Nov.
Draconarius tryblionatus (Wang and Zhu, 1991), NEw COMBINATION (from Coelotes)
Draconarius uncinatus (Wang et al., 1990), NEW COMBINATION (from Coelotes)
Draconarius yichengensis, sp. nov.
Key To Females of the Species of the Genus Draconarius
wmipicynal teeth present... 5/55 4 0s eer. sw ks 3) sa dl tee 2
Epigynal teeth absent
ynal teeth long, with length at least five times width (Fig. 8A) ..............--.--- 3
ynal teeth short, with length at most four times width (Figs. 19A; 39A) ............. 3)
ynaliteethi with apexes close togethen (Fig. 8A): =. .422¢. 4-54 4255e eee acidentatus
ynall\teeth with apexes widely separated ....... 455.2. 552082))2202 =e eee =
WANG: EAST ASIA COELOTINE SPIDERS S11
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mm Spemmathecae withiantenonendsyclose itogethemisarcis s.s5). Secs lat eee huizhuensis
Spermathecac with anterior ends widely separated) 5... ..2 eae strophadatus
. Epigynal teeth with bases close together, separated by less than their width (Fig. 26A)
RP RS) ered gens, arth Wy eee enact ul eS oye AE oR SUD SA digitusiformis
Epigynal teeth with bases separated at least by their width (Figs. 13A; 24A)............. 6
. Copulatory ducts expanded anteriorly anterad of spermathecae........................ a
Sopulatory ucts Otherwise HE LEGA aS se OTS LN SAREE, ARIAT ha PON UEARS Et AS 14
. Epigynal teeth situated posteriorly near epigastric furrow ...................... infulatus
Epigynal teeth situated anteriorly, widely separated from epigastric furrow .............. 8
. Epigynal hoods deep, situated laterad of atrium (Figs. 56A; 58A)...................... 9
Epigynal hoods shallow, situated anterad of atrium (Figs. 9A; 44A)..................0. 10
. Spermathecal bases widely separated, stalks long and looped; spermathecal heads situated
anichadOisspeninathecac (B19 DONS) ioe. are eawe Perepenny ete ace ats aay UNI tr atten aig cael rotundus
Spermathecal bases close together, stalks short, not looped; spermathecal heads situated laterad
OMSpeninathecde (EIS? DO) seta ccs cs sre rel see conte eee RE a eras ee fame oh schenkeli
Epreynaliteeth situated near atrial mareimi(Rigs 9A SOA) ein crite «hirer sire. ae 11
Epigynal teeth widely separated from atrial margin (Fig. 44A)....................04. 12
. Spermathecal stalks laterally extending; spermathecal heads surrounded by looped copulatory
eer ese (Estes OB) isa ey xray veh tr eos ae oo Seana hie Pag ehear sie, oh SEND in cota as ia, Slo aya Yoav ee gts penicillatus
Spermathecal stalks short, broad, not laterally extending; spermathecal heads not surrounded
by .copulatory-ducts (Rigs: OB iris) | sete va tpta shears tae eee ashe Renee adligansus
. Epigynal teeth separated by less than half atrial width (Fig. 44A)............ neixiangensis
Epigynal teeth separated by at lease atrial width (Figs. 25A; 34A).................... 13
mSpeLiMathecal heads snort. postenory situated (Rig. 25B) cia cee sc. oe ins oe denisi
Spermathecal heads long, anteriorly situated (Fig. 34B) .................. hangzhouensis
Spermathecae short, rounded, with length almost same as width (Figs. 43B; 32B)....... HS)
Spermathecae elongated, with length at least twice width (Figs. 61B; 67B; 39B) ........ Dy
mopermatnecal heads not visible trom: dorsaliview (Fig: 3ZB)\e2 0. eats ks Sissies ate aece 16
Spermathecal heads visible from dorsal view (Figs. 30B; 43B) ..................004. 7
. Copulatory ducts broad, situated mesad of spermathecae (Fig. 60B)............ singulatus
Coapulatory ducts not visible from dorsal view (Fie, 32B) 2. ee se ee eis gurkha
. Spermathecal heads situated anterad of spermathecae (Fig. 69B)..................4.. 18
Spermathecal heads situated mesad of spermathecae (Figs. 30B; 43B)................. Zi
. Epigynal teeth long, with length at least twice width; spermathecal heads situated
AMUSKOLALE Ala (ENS OOS) meryaryreety tea cers ate ene cae carlo -4, oes f tleraonedae te pane yichengensis
Epigynal teeth short, with length at most same as width; spermathecal heads situated
LIT CPAa 6) yg ee hee aero py rarer en ike eh Sa De RLM EE AL CSSA ae LER Ca a A Ma id aco UE hac I)
Epicvnal tecth separateaeiromn atiimiyy.tyn) atv ays io heer tics tant tetas Faucets himalayaensis
Episviral tecth situatedmmeatiatriuini ee, trees 2G Aad ici Pe Lanta taint tame ee nia erste apts hooceebebet ween: 20
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Spemmathecacicloseitogethers... 2... 2. : See Se Bie eee ee eee altissimus
Spemmathecae slighthysseparated!.”. .).-). be coe een sree Serene cei eee subtitanus
. Epigynal teeth widely separated; copulatory ducts and spermathecal heads situated mesad of
Spermathecac(RissMs OASB) i 2s tS Jie tthe se te heen ee RO ae ae ae coe are episomos
Epigynal teeth slightly separated; copulatory ducts originating dorsally and spermathecal heads
originating ventrally on spermathecae (Figs. 43A—-B).....................--- nanyuensis
Spermathecae broad, anteriorly expanded; spermathecal heads situated medially on
Spenmathecae (Eves: ISB 61B267B). 22.6.6: ne Bee ae 1 ee DB
Spermmathecaciotherwise eee es ee oe eee eet et hove oe ce et 40
Epigynal teeth situated anteriorly, close together (Figs. 17A; 24A)..................-- 24
Epigynaleteeth widely separated'(Figs: I3A;/67 A)... 25. 2. 22 ne ee ee 26
AMtenon atrialumarcin broad, biturcate (Pigs 173) 2k. a aes calcariformis
Anterior atrial maroinindistinct, not bifurcate 4422.25. 925% oe ee ee 25
Spermathecaluheads indistinct (Fis. 2418) 2 a cee ae oe ee davidi
Spermathecall headsidistinct.,< -/.5 1. asc awices erclacieh eis el oer ee colubrinus
Epigynal teeth wide apart, at least 1.5 times atrial width (Fig. 67A)................... Dah
Epigynal teeth separated by atrial width or less (Fig. 3A) .... 22. 1.25.4 ee Sil
Spermathecal stalks with anterior extension not converging..................-- yosilanus
Spermathecal stalks anteriorly converging, close together (Fig. 22B).................. 28
Epigynal' teeth situated! posteriorlysnear atrium’ 22% 9). 5: . 82s a ee picta
Epigynal teeth situated anteriorly, widely separated from atrium (Fig. 22A) ............ 29
. Epigynum with a broad, membranous, transverse extension in front of atrium (Figs. 22A—B)
SOMO Ooo DOD oO oO OO OD Om Oyo Golo) oldOy0 010 0) Choco 01d 0.0 00 dia M0) O10Nd 0 0-0W1d CIB O00.0 6.0.0 0.0 coreanus
Spermathecal heads situated on anterior 1/3 part of spermathecae; spermathecal bases broad
(Ei OB) acta coven. sicpe tory veasatt sake ecole hecho eat ee eee he eee wudangensis
Spermathecal heads situated on posterior 1/3 part of spermathecae; spermathecal bases narrow
SMR STR ENCE «Sab. nanictbets Senki as Mater «AY aioe 4 RO Pod woe ORL oO Shee EN a venustus
. Epigynal teeth situated posteriorly, at same level or posterior compared to anterior atrial
HUN AT OUT pe rcs Pah ah sre ote chan 3S Serene SPOS: hfe en SGGN she RUS SUIS ARI ae tT GT Seg eae 32
Epigynal teeth situated anteriorly compared to anterior atrial margin (Fig. 13A)......... 35
ss penmathecalibasesiclose together... ; ..45...6 ee eee eee funiushanensis
Spermathecal bases separated at least by their- width J... 22 5.2 2252 224p ee eee 33
. Epigynal teeth situated at same level as anterior atrial margin.................-.-.-- hui
Epigynal teeth situated at posterior level compared to anterior atrial margin............ 34
. Spermathecal heads situated medially on spermathecae .................... gingzangensis
Spermathecal heads situated on anterior 1/3 of spermathecae.................. linzhiensis
. Spermathecae with anterior expansion broader than stalks (Fig. 13B) ................. 36
Spermathecae with anterior expansion same or narrower than stalks (Fig. 61B) ......... 38
WANG: EAST ASIA COELOTINE SPIDERS 513
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46.
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Epigynal teeth broad, situated slightly anterior of atrium .................... trifasciatus
Epiaynal tecth narrow, widely separated fromvatrium 5 ).)544 42 ise. Oe) oe en. Si)
. Anterior atrial margin expanded posteriorly (Fig.13A).................00.0005 aspinatus
AmienOn atrial marcin notiexpanded postemorly.......60+>4-5-50-+0500045- gyriniformis
. Copulatory ducts originating medially, looped laterally (Fig. ISB) .......... baxiantaiensis
Copulatory ducts originating medially, short, not looped (Fig. 61B)................... a9
. Spermathecae widely separated, at least by their width (Fig. 61B).............. stemmleri
SHemnathecae Situated Close together ss sas ania tens eer nea Gyan deen ee eaer ae yadongensis
Spermathecal stalks long, slender, extending laterally, and converging anteriorly (Fig. 39B)
co Ge SNS I ena OS OUR NE eee Penna OOP n RON Orsarts.| SUMP SOC Gnue Porat ee eae ean API or 4]
Spemmathecal stalks) broad (Figs. 19B. 5B) iene. Caen rans sf oueeechch weyers noe ousie. 47
. Copulatory ducts extending along spermathecae, not looped (Fig. 21A)................ 42
Copulatory ducts looped. around: spermathecae. (Fig, 39B). 22x. yee ee eo es ee ee 43
. Epigynal teeth situated laterally, widely separated; spermathecal bases broad, heads long, close
POP UMC Ie (RIGS SOA ES) oa cee tlateclataithede Peay: SemAE Oooo Sette aie ete ac ers dee easieuer Jiangyongensis
Epigynal teeth situated medially, moderately separated; spermathecal bases narrow, heads
midehwsenaraed (Figs: 21A—B)k- sts aewith ida aes tl ented es a bees cheni
BE pioymal teeth situated anteriorly, close together (Fig: 66A) 7... 2.2... 2.662. 5scc5ene8: 44
Epigynal teeth situated near atrium, widely separated (Fig. 39A)................00005 45
. Spermathecae with anterior extension extending anteriorly; spermathecal heads not covered by
capelatonyducts:(EIe x60B) ssy.4 asa ae AO Eek Se Ee 2 wenzhouensis
Spermathecae with anterior extension extending medially; spermathecal heads covered by cop-
EH ORYECUCtSa(h1 93155) ))x-gyras PSAP, seb Alaetss et le he EEN Suedab ous tees: sched Litera labiatus
Epigynal teeth separated by less than half atrial width (Fig. 11B)................ arcuatus
Epipynal teeth separated by at least atrial width (Fig. 389A)» 2. 22. ei een eee 46
Epigynal teeth situated laterad of atrium, away from atrial margins; copulatory ducts looped
mMearallyzon spenmmathecac, (MisshOSA—B)ist) wets keine) em eet arucesn .eaedenle wuermlil
Epigynal teeth situated near atrial margins; copulatory ducts looped anteriorly on spermathecae
(TET Be, Shoda Sl BS) hab a dy te oe ge chien eta MOR at eins cS RE ROE ea a SPR A a lutulentus
Copulatory ducts looped around spermathecae (Figs. 19B;51B) ..................... 48
SOpuUlaronyrcuctsmOPlOOpPed s AA ee eae he ON oe VEN OHA Ee Great Soar 2 49
Copulatory ducts originating medially, with two loops around spermathecae (Fig. 51B)
MMP Wet Mel ce eh cn neta Mabie eB) Shc Ney galls AR hte tn Re COLE HOE THAD SCRE potanini
Copulatory ducts originating laterally, with one loop (Fig. 19B)................ carinatus
Spemiuathecal bases with diverticula (Fig. 48B))....2.-.-42: 2-212 s555ee os paraterebratus
Spemnatiecadl basesswithout diverticula... ce eee an See ee ee es See 50
Pe petmMAtecal bases situatedieloseditoeethersn 22 5926 500. 2 dela ono eee pervicax
Spermathecal bases separated at least by their width (Fig. 55B)...................... 51
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Epigynal hoods situated posteriorly near epigastric furrow; copulatory ducts large; spermathe-
Calestalks) broads not convoluted (E1gs) 55 A—B)=5 see eee eee quadratus
Epigynal hoods situated anteriorly; copulatory ducts large; spermathecal stalks convoluted
Gio sap AASB) Bene oe eerie. ce © Foe wk wane ccle eeaerd loner aes seeker Pay eee pseudowuermlii
Copulatory ducts originating mesad of spermathecae, extending laterad of spermathecae with
A= OOPS (EIS OTB) s cece te oe cine nos kee oot tee 6 ce aa Se Ae eee rufulus
Copulatory ducts originating either mesad or laterad of spermathecae, not extending laterad of
Spemmathecaeand looped. = % 5, <2 Pes pics che yore osahs =, heey win) G/e. ch che) ehoaegs 9) Se eee 53
Copulatory ducts situated laterad of spermathecae (Fig. 31B)...................-.... 54
Copulatory ducts situated mesad of spermathecae (Fig. 33B)....- -. 3-402 pee eee 56
Copulatory ducts only laterally extending, not anteriorly expanding; spermathecae anteriorly
converging» notlooped (Fig: 23B)). sn «ees: 2 noes ee ee eee griswoldi
Copulatory ducts expanding anteriorly; spermathecae looped (Fig. 23B)............... a5)
Atrium with distinct lateral margins; epigynal hoods situated near atrium; spermathecae looped
laterallya(Big= AB) eves. 3 tgs thon bole cpu pack ea eee ee absentis
Atrium with distinct anterior margin; epigynal hoods widely separated from atrium; spermath-
ecae looped medially (Fis: 23A—B) ooo cle wc, desu oth ote cee eicie tae kt eee Curlosus
Epigynal hoods situated laterally, laterad of atrium (Fig. 42A)....................25. S//
Epigynal hoods situated anteriorly, anterad of atrium (Fig. 27A) ...................-. 3)
Copulatory ducts notlooped (Fie 63B)R as. 252 ee eo eee eee terebratus
Copulatory ducts looped around spermathecae (Fig. 42B).......:....52. 4-50 58
Copulatory ducts with two loops; spermathecae with distal end strongly convoluted (Fig. 42B)
BNE SAN Stee ATS Be Ree a wna Succ CE ER, See OS aoe dar oo NSE reEtohh SARE eEne molluscus
Copulatory ducts with one loop; spermathecae with distal end smooth, not convoluted (Fig.
5 011 }) i ara ne ae ieee ORS woven eae te Cre AE ty NN MN RRR Sec 6 a0 haopingensis
Spermathecae elongated; looped >... 102 205. 2 sas Sas 0. Dee a 60
Spermathecae short, not looped . 2... 6... 0s oss ee ee 2 oe ok eee 62
. Spermathecal bases extending laterally, away from each other ............... tryblionatus
Spermathecal bases extending medially, approaching each other ..................-.- 61
. Spermathecal stalks strongly convoluted, shaped like circles.................. syzygiatus
Spermathecal stalks extending laterally, then converging medially, not shaped like circles
Bp Rete ausi ee (5/ syeiielavieia eile aed. s Soeia tls Se wie de Soe we ee ee streptus
2. Copulatory ducts broad, strongly extending anteriorly (Fig. 41B).............. magniceps
€opulatory ducts small, not anteriorly extending...........-++..+5---+ epee 63
. Epigynum wrinkly anterad of atrium as in Figs. 27A;47A.............2.0000000000- 64
Episynummot wrinkly;..... ...c¢ be... .<0Sh i808: Glace ee BER See 67
Spermathecal bases broader than stalks (Fig. 47B) .....2.5....254. --5 2 eee 65
Spermathecal bases narrower than stalks (Fig. 27B) .............-.-.-7, > 5eeeeeeeee 66
Spermathecal heads situated anteriorly on spermathecae (Fig. 47B).......... parabrunneus
Spermathecal heads situated medially on spermathecae (Fig. 52B) ........ pseudobrunneus
WANG: EAST ASIA COELOTINE SPIDERS StS)
SeeSpckMatmecae withiconvergina distalvemdsi; sacs cele sie steers ciciene cleus late dee eee brunneus
Spermathecae with widely separated distal ends (Fig. 27B).................... disgregus
67. Spermathecae strongly expanded anteriorly; spermathecal heads situated medially on sper-
TAU ROCRS (S64 3) Rae cate tats Geen ee een EL CTEM Cry anna Teer en, ated striolatus
Spermathecae not anteiorly expanded, or slightly expanded; spermathecal heads situated
ABICIEORYZONES PETIMALNE CACY (Ue12s WOES) see exe cierto cies See ene ccs ee te cae ate rsa ee ane ee 68
peESecemathecac medially: lobed (Eig: 3B) sau. tos cere: hese coors) sinter oe aye Sas Soa ore 69
CRIM ALME CACO’ LODEC cs ire we cian acs ee bye Sasucy eegel cM eee yoeG eM Cos TAN icant tare eka it 70
69. Spermathecae with lobes close together, slightly separated (Fig. 18B).......... capitulatus
Spermathecae with lobes separated at least by their sizes (Fig. 53B)...... pseudocapitulatus
70. Spermathecae widely separated at least by their sizes (Fig. 28B)..................... Di
SpeEmathecac separated py less than) them sizes (Figs 29B) trae oe ices oe ee oe 12
71. Spermathecal heads and copulatory ducts distinct (Fig. 28B) ....................- dissitus
Spermathecal heads and copulatory ducts indistinct..................... amygdaliformis
ie spermathecal bases broader than stalks (Fig. 29B)....5..2-5.2 5222552256 - seer se dubius
SpcemiathecalgbasesmManrOw.em thant Stal KSesrusit a: cies emo tor ete Sateen) tiers ise eee isa 1
73. Copulatory ducts large, anteriorly extending; spermathecal heads extending mesad of sper-
MIAME CASKET OO OES) ore cs crave lars Matrstacct ice Saimen aie Heys Anaheim aloes PN Boe OPE simplicidens
Copulatory ducts small, not anteriorly extending; spermathecal heads extending anterad of
Sem Abneedes (10s AOR iiss cys. uck teens core eae a ote eee ae eae eee patellabifidus
Key To Males of the Species of the Genus Draconarius
Peat lanapopnysis‘absent (Big. ISB)". oo. s arcs aoe ee cease A eee ee eR ee een te 2
Pate tim apoOply sis present (Bip+39 Brae es eee tee atue ep ct he ene oie eo eae hed eet tee ae heg e 12
Pam iceHat apophysis-not spoon-like: (Rigs: 19C-B) iin oc ger ee Sele ae bes ae on ees 3
Median apopuysisspoonziike (Bic. 13D) xc... caaschaye nae paperehteal adeno wee ue Siemon ee 4
Median apophysis strongly bifurcate (Figs: 19C-B)2 32. 2.2.22..-.2. 2205). 2.-5e carinatus
Rie diamapopuysis,cloneated, not biturcate. 4-455 1 oo erie Acne ae acti nudulus
4. Conductor long, extending posteriorly, reaching embolic base (Figs. 17B; 57D) .......... 5)
Conductor short, retrolaterally or anteriorly extending (Figs. 13D; 44C)................. 6
MEE IbOls Dioad posterior imionsin,| (Figs. S/E.D)ins .. 2.2hN. te eee ee eee rufulus
Embolus slender, retrolateral in origin, (Figs. 17A-B)..................-.. bituberculatus
PeEmvolns protateral mm orgim (Fig. TIAN ac, en oe wiz elec cee neo eue 2 ocr age one teal eee 7
Pambolus posterior invoriein (F1e. U3) i. norte gaycts cicipeyo oye ors) toned sins See ioe 8
7. Conductor with slender apex; median apophysis strongly elongated (Fig. 12)...... argenteus
Conductor with broad apex; median apophysis slightly elongated (Fig. 20) .. . . chaigiaoensis
8. Embolus broad, conductor broad, deeply grooved (Figs. 38A, 44C) .................-.. 9
Embolus slender, conductor slender, slightly grooved (Figs. 13C-E)................... 11
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. Embolus with narrow base; median apophysis not elongated (Figs. 38A—B)...... linxiaensis
Embolus with broad base, median apophysis strongly elongated (Fig. 44C)............. 10
Embolusswathislenderapexeie. cee len foes. no Boece eee ens ot en ee colubrinus
Embolus;wathibroadiapex (Bigs: 44€ ID) 23. v2 -is5e er aa oe eee neixiangensis
Conductor with dorsal edge slightly toothed, as in Figures 8-43:3—5 (Hu, 2001). . . altissimus
Conductornot toothed (Figs. 13GB) 9.0 2. Se a ee eee aspinatus
Ratellar‘apophysis bifurcate (Fig 49D) 2206.03. so ge. eo eee 113)
Patellarapophysis not bifurcate/(Figs: 39D). 2 8. 208) | 02) a eee 16
Medianvapophysis spoonzlike (Fig. 65) 2350. 3): 3 45 ee eee uncinatus
Median apophysis not spoon-like (Figs. 49€, D).2 05. ...503...) eee 14
Conductor with distal part spiraled) ye... 2 ae oe ee eee tryblionatus
Conductor not spiraled... oo. 2s. So ei 5 ee ee te oe IS
FE mboliG.DaSeMOtCME Gas .25..64.. cate hue cue cutee snas Val ago ap ae himalayaensis
Embolic base mot notched (igs. 49C Da ean pees Ne eee ee patellabifidus
Median apophysis absent icc. 2°. /40..60 see we es ee ae ge oe ee. ee 7
Median.apophysis present .6.25.. i426 6 stipes bans beige 6 Sas ss ee 18
Embolus broad (Fis: 23€) oc ft os racine oe pl ee aac each eee Curiosus
Embolus:slender (Bigi 7C)is.n 6.008 tee oust ace « Soles us aye's eos SE oe absentis
Median apophysis not spoon-like:(Figs. 27€; 29C) 0.2. 5. ee. ae oe ee 19
Median. apophysis spoon-like (Fig. 67D)... «2... aon Wee oboe ee | eee zal
Median apophysis broad; patellar apophysis strong curved (Figs. 27C, D)........ disgregus
Median apophysis slender; patellar apophysis not curved (Fig. 29C) ................-- 20
. Patellar apophysis as long as or longer than tibia (Figs. 29 C, D).................. dubius
Patellar apophysis much shorter than tibia (Figs. 33A-B)....................4-- incertus
. Conductor as long as cymbium, looping and extending posteriorly ............. syzygiatus
Conductor SHOT. 5c: cag So aces woo 6 the bis PORE ol crs EL lone core Si eee eee 22
wEmbolicibase lobed (Fig.63D)* 2220.8 i225. ee etc ole ae eee terebratus
Embolic:basenot lobed |. ).icc,.,., . «,s s.acs.0,0,5 <2 /snegele, «0. 5 oa, nasvacustegeneate, Seinen eC Wr)
Conductor strongly bifurcate (Figs. 16C€;46B) 2.) 02.57.3525 o) oe 24
Conductor not: bifurcate. oo. i... sc 6 he cums dis ae oeee nde eer eee 26
Conductordeeply bifurcate from the base (Fis=16@)) 45-2.) eee calcariformis
Conductor slightly bifurcate from the distal part (Fig. 46B) ..................+..--- 25
25. Embolus broad, cymbial furrow short (Figs. 46A—B)"....2. 5.2202) ee eee ornatus
Eimbolus’siender cymbialturrow long...) 2s.) ee ee ee huizhunesis
. Conductor with apex slightly spiraled, anteriorly extending (Figs. 26C-E)..... digitusiformis
@onductor slender, retrolaterally extending (Fig. 67D)... 2.22 27
WANG: EAST ASIA COELOTINE SPIDERS 517
Meenate lan apaphysisiasi Ongvasitiblayas aajee. Cieoctcas Saas aerate ae funiushanensis
Batelarapaphysisishor= less thanitibralalene thier tae ataek nee eran nie nee Seda ee 28
PE MoOIs prolatcralimeon oun (EIS. OOD) a auc eepewho nap nai eews cack cy None eo aati 29
Fanboluciposten Onin Omeimy (1 OW DD) ces coe cee as eae ee Bie eek ee cy eee ls egy ek 30
PeeWeAwlonger than half tibial lensth\(Fis- G0E)- 225... 55. -+. ee eee eee singulatus
RWeawshorter tian halttbialllensth? Sooo Soe ee wee ens ye ae ee nD Meee everest
See Acshortemthan half tbiallength(Figs.39D iB) ins. soe aceite Sera, tens eee ee 31
Reson scrmthanshaligubialylensthy(Figs. 22D, E)eiciesen cen see ore ene tne eae 32
eonauctor lone, looped posteriorly, (Fig.39D). scsi. seo es nese ene ae lutulentus
WanGuctomsnon not loopedh(EIG GMD). 5 seca ae oes eee oe ee wudangensis
32. Median apophysis not extending, with anterior edge not free (Figs. 22D; 50D).......... 33
Median apophysis extending, with anterior free edge (Figs. 1OA; 31C)................ 36
Sa eymbial furrow-less, than half cymbial length (Pig. J4C) 2.22.45 cn bes. dn ee 34
Sumpialiunrowsat least halacymbialilenethy(Big.22©) a2 a foe cece aie oa eae ae 35
Sa@onductor with broad anterior'edge (Fig: 64B):. 00.522. en ie ees tibetensis
Conductorwithout broad anterior edge (Fig. TAB) io. es i en ee sel ee baronii
35. Embolic base short, conductor positioned posteriorly (Fig. 50D).............. penicillatus
Embolic base long, conductor positioned anteriorly (Fig. 22D) ................. coreanus
PE MESO TCAD ASE NSMOL a, rhe ree eee ree nt ere Oe A eR EEO ee eee venustus
paibolieanascreoncated (Eton SiC) re see perce ee eee ene eee nl te eee ae Sy
37. Conductor with narrow dosal edge; embolic base with a small lobe on prolateral side
ET Fee. Stas) RIA coh cs SEA Sd Sica ou TS SEEN RASTER VR AREAS hates sted trifasciatus
Conductor with broad dosal edge; embolic base normal, without lobe (Figs. IOA-B) .... 38
RemmEDEEERSOLTI SE SCOU E(t Doel VAN) eas sae, ebay Aon) Seek cicL ais cates eRe ee ie tas oS Soe SR agrestis
Bribolusrslendernm (Bie Si©) ake nits ot Sie eg Pa ed ieee eee Se a tthe griswoldi
Draconarius absentis Wang, sp. nov.
Figures 7A—D; Map 3
Typres.— Male holotype and female paratype from Luoshuidong, 28 air km E TengChong,
24°57'N, 98°45’E, native forest, 2300m, Baoshan Prefecture, Yunnan, China (October 26-31,
1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI; 9 female and 7 male paratypes
from Pass over Gaoligongshan at 2100 m, Nankang, 36 air km SE TengChong, 24°50/N, 98°47’E,
native forest, Baoshan Prefecture, Yunnan, China (November 4—7, 1998; C. Griswold, D.
Kavanaugh, C-L. Long), deposited in HBI (7 females and 5 males) and CAS (2 males and 2
females); 3 male and 5 female paratypes from Luoshuidong, Baoshan, Yunnan, China (October
26-31, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in CAS.
ETYMOLOGY.— The specific name refers to the absence of both epigynal teeth and median
apophysis, which differs from most other Draconarius.
DIAGNOSIS.— This new species can be distinguished from other Draconarius by the absence
518 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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of epigynal teeth, the broad copulatory ducts, the short spermathecae of female (Figs. 7A—B), and
by the short cymbial furrow, the trifid conductor, prolateral origin of embolus, and the absence of
median apophysis of male (Figs. 7C—D).
FEMALE.— Total length 9.80. Carapace 4.60 long, 2.90 wide. Abdomen 5.20 long, 2.20 wide.
Eye sizes and interdistances: AME 0.11, ALE 0.20, PME 0.18, PLE 0.20; AME-AME 0.10, AME-
ALE 0.07, PME-PME 0.15, PME-PLE 0.20, AME-PME 0.17. Leg measurements: I: 12.0 (3.40,
4110; 2:80, 1.80); I: 10.8 (G-00; 3-50, 2:60, 1.70); I: 9:76 (2.60, 3:16; 2:505 1°50) IWS wAG0)
4.30, 3.60, 1.90). Chelicerae with three promarginal and two retromarginal teeth. Epigynal teeth
absent; atrium large, with broad anterior atrial margin; epigynal hoods deep, situated near anterior
atrial margin; copulatory ducts broad, extending anteriorly and laterally; spermathecal heads large,
situated laterad of spermathecae; spermathecal bases small, widely separated; spermathecal stalks
separated, anteriorly extending and diverging (Figs. 7A—B).
MALE.— Total length 9.60. Carapace 4.80 long, 3.10 wide. Abdomen 4.80 long, 2.80 wide.
Eye sizes and interdistances: AME 0.16, ALE 0.21, PME 0.19, PLE 0.20; AME-AME 0.08, AME-
ALE 0.06, PME-PME 0.10, PME-PLE 0.19, AME-PME 0.16. Leg measurements: I: 14.3 (3.80,
4.86, 3.50, 2.10); II: 12.6 (3.40, 4.12, 3.10, 2.00); III: 10.6 (3.00, 3.60, 2.20, 1.76); IV: 15.4 (3.90,
4.80, 4.50, 2.20). Chelicerae with three promarginal and two retromarginal teeth. Male palp with
patellar apophysis short; RTA long; lateral tibial apophysis large, near RTA; cymbial furrow short;
conductor moderately long, with trifid apex; conductor dorsal apophysis present; conductor lamel-
la small; embolus short, prolateral in origin; median apophysis absent (Figs. 7C—D).
DISTRIBUTION.— China (Yunnan) (Map 3).
OTHER MATERIAL EXAMINED.— None.
Draconarius acidentatus (Peng and Yin, 1998), NEW COMBINATION
Figures 8A—B; 97A; Map 3
Coelotes acidentatus Peng and Yin, 1998:26, figs. 1-3 (female holotype and | female paratype from
Huangsang, Suining, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:365.
DIAGNOsIs.— The female of this species can be easily recognized by the long, closely situat-
ed epigynal teeth, the deep, medially situated epigynal hoods, the laterally originating copulatory
ducts, and the posteriorly situated spermathecal heads (Figs. 8A—B).
DESCRIPTION.— Described by Peng and Yin (1998). Chelicerae with three promarginal and
two retromarginal teeth. Female epigynal teeth long, broad, originating anteriorly, situated close
together; epigynal hoods deep, situated medially; atrium small, near epigastric furrow; copulatory
ducts originating laterad of spermathecae, anteriorly extending; spermathecal heads long, originat-
ing posteriorly laterad of spermathecae; spermathecal bases broad, slightly separated; spermathe-
cal stalks broad, close together (Figs. 8A—B). Male unknown.
DISTRIBUTION.— China (Hunan) (Map 3).
MATERIAL EXAMINED.— CHINA: Hunan: Suining, Huangsang, August 1996, female holotype
(Yin, C.M. and X.J. Peng, HBI); Suining, Huangsang, May 28, 1996, 1 female paratype (Yin, C.M.,
X.J. Peng and Y.J. Zhang, HBI).
Draconarius adligansus (Peng and Yin, 1998), NEW COMBINATION
Figures 9A—B; 97B; Map 3
Coelotes adligansus Peng and Yin, 1998: 26, figs. 4-6 (female holotype from Nanyue, Hunan, China, in HBI,
examined).
WANG: EAST ASIA COELOTINE SPIDERS 519
DiaGNosis.— The female of this species is similar to D. denisi in having the anteriorly
expanding copulatory ducts but can be distinguished by the posteriorly situated, moderately sepa-
rated epigynal teeth (situated near atrium, separated by less than atrial width) and the anteriorly
expanded spermathecae (Figs. 9A—B).
DESCRIPTION.— Described by Peng and Yin (1998). Chelicerae with three promarginal and
two retromarginal teeth. Female epigynal teeth short, situated on anterior atrial margin, separated
by less than atrial width; atrium with distinct median carina; copulatory ducts large, anteriorly
extending; spermathecal heads small, anteriorly situated; spermathecal bases small, widely sepa-
rated; spermathecal stalks short, anteriorly expanding and slightly diverging (Figs. 9A—B). Male
unknown.
DISTRIBUTION.— China (Hunan) (Map 3).
MATERIAL EXAMINED.— CHINA: Hunan: Nanyue, August 3—7, 1995, female holotype (C.M.
Yin, HBD).
Draconarius agrestis Wang, sp. nov.
Figures 1OA—B; Map 4
Types.— Male holotype and 2 male paratypes from from Luoshuidong, 28 air km E
TengChong, 24°57’N, 98°45’E, native forest, 2300m, Baoshan Prefecture, Yunnan, China (October
26-31, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (holotype male and 1
male paratype) and CAS (1 male paratype).
ETYMOLOGY.— The specific name refers to the large cymbial furrow and long, broad embo-
lus.
DIAGNOSIS.— The male of this new species is similar to D. griswoldi in having a small patel-
lar apophysis, an elongated cymbial furrow, and a posteriorly originated, long embolus but can be
distinguished by the broad embolus, the broad conductor apex, and the spoon-like median apoph-
ysis (Figs. 1OA—B).
MALE.— Total length 9.60. Carapace 4.80 long, 3.40 wide. Abdomen 4.80 long, 4.00 wide.
Eye sizes and interdistances: AME 0.16, ALE 0.20, PME 0.21, PLE 0.21; AME-AME 0.09, AME-
ALE 0.05, PME-PME 0.09, PME-PLE 0.24, AME-PME 0.11. Leg measurements: I: 13.7 (3.92,
4.60, 3.20, 1.94); Il: 12.1 (3.40, 3.92, 2.80, 1.80); Il: 8.18 (2.60, 1.92, 2.36, 1.30); IV: 12.4 (3.50,
4.24, 3.20, 1.50). Chelicerae with three promarginal and two retromarginal teeth. Male palp with
small patellar apophysis; RTA long; lateral tibial apophysis small, widely separated with RTA;
cymbial furrow large, longer than half cymbial length; conductor broad, with broad, membranous
dorsal edge and large basal lamella; conductor dorsal apophysis present; embolus posterior in ori-
gin, long, broad; median apophysis spoon-like, elongated (Figs. 1OA—B).
FEMALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 4).
OTHER MATERIAL EXAMINED.— None.
Draconarius altissimus (Hu, 2001), NEW COMBINATION
Map 3
Coelotes altissimus Hu, 2001:131, figs. 843:1-5 (female holotype and male paratype from GongbuJiangda,
Tibet, China, in SDU, not examined).
DiAGNOsIS.— The male of this species is similar to D. aspinatus and can only be distinguished
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by the slightly toothed conductor. The female is similar to D. himalayaensis and D. subtitanus in
having the short epigynal teeth, anteriorly situated spermathecal heads and rounded, closely situat-
ed spermathecae but can be distinguished from D. himalayaensis by the epigynal teeth position (sit-
uated close to atrium), from D. subtitanus by the non separated spermathecae.
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal and two retromarginal
teeth. Female epigynal teeth short, widely separated, situated near anterior atrium; atrium small,
posteriorly situated; spermathecal heads situated anteriorly; spermathecal bases widely separated;
spermathecal stalks broad, rounded, close together. Male palp without patellar apophysis; RTA
long; lateral tibial apophysis small; cymbial furrow large, longer than half cymbial length; conduc-
tor slender; embolus long, posterior in origin; median apophysis spoon-like, elongated.
DISTRIBUTION.— China (Tibet) (Map 3).
MATERIAL EXAMINED.— None.
Draconarius amygdaliformis (Zhu and Wang, 1991), NEw COMBINATION
Map 4
Coelotes amygdaliformis Zhu and Wang, 1991:2, figs. 8—9 (female holotype, 2 female paratypes from Xishan,
Kunming, Yunnan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:365, figs. 216H-I.
DIAGNOSIS.— The female of this species is similar to Tegenaria domestica by having small,
posteriorly situated atrium and small, widely separately spermathecae.
DESCRIPTION.— See Zhu and Wang (1991). Chelicerae with three promarginal and two retro-
marginal teeth. Female without epigynal teeth; atrium small, posteriorly situated, near epigastric
furrow; spermathecae small, rounded, widely separated. Male unknown.
DISTRIBUTION.— China (Yunnan) (Map 4).
MATERIAL EXAMINED.— None.
Draconarius arcuatus (Chen, 1984)
Figures 11A—B; Map 4
Coelotes arcuatus Chen, 1984:2, figs. 34 (4 female paratypes from Huanglongdong, Hangzhou, Zhejiang,
China, in HTC, examined).— Chen and Zhang, 1991:189, figs. 187.1-2;— Song, Zhu and Chen,
1999:374, figs. 216J—K, 218A-B.
Draconarius arcuatus: Wang, 2002:66.
DIAGNOsISs.— The female of this species is similar to D. cheni in having a broad, medially sit-
uated, anteriorly expanding copulatory ducts but can be distinguished by the broad atrium (twice
epigynal teeth distance) and the distinct, anteriorly situated spermathecal heads (Figs. 11A—B).
DESCRIPTION.— The female was described by Chen (1984). Chelicerae with three promargin-
al and two retromarginal teeth. Female epigynal teeth situated posteriorly near atrium, close togeth-
er; atrium slightly extending posteriorly; copulatory ducts originating posteriorly, extending ante-
riorly, slightly spiraled; spermathecal heads distinct, anteriorly situated; spermathecal bases small,
widely separated; spermathecal stalks long, laterally extending (Figs. 11A—B). Male unknown.
DISTRIBUTION.— China (Zhejiang) (Map 4).
MATERIAL EXAMINED.— CHINA: Zhejiang: Hangzhou, Huanglongdong, February 26 to March
1, 1982, 4 female paratypes (Z. F. Chen, HTC).
WANG: EAST ASIA COELOTINE SPIDERS 521
Draconarius argenteus (Wang et al., 1990), NEW COMBINATION
Figures 12A—B: Map 4
Coelotes argenteus Wang et al., 1990:229, figs. 117-119 (male holotype from Jinhong, Yunnan, China, in
HBI, examined). — Song, Zhu and Chen, 1999:374, figs. 218C, J.
DriAGNosis.— The male is similar to D. nudulus in lacking a patellar apophysis and having a
short conductor but can be distinguished by the short RTA (half tibial length), the spoon-like medi-
an apophysis, and the lobed embolic base (Figs. 12A—B).
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and five
retromarginal teeth. Male palp without patellar apophysis; RTA approximately half tibial length;
lateral tibial apophysis large, situated anteriorly and near RTA; cymbial furrow slightly less than
half cymbial length; conductor broad, with slender apex; conductor dorsal apophysis present; con-
ductor lamella broad; embolus prolateral in origin; embolic base with a lobe on its retrolateral side;
median apophysis spoon-like, strongly elongated (Figs. 12A—B). Female unknown.
DISTRIBUTION.— China (Yunnan) (Map 4).
MATERIAL EXAMINED. — CHINA: Yunnan: Jinhong, October 21, 1987, male holotype (J.F.
Wang, HBI).
Draconarius aspinatus (Wang et al., 1990)
Figures 13A—E; Map 5
Coelotes aspinatus Wang et al., 1990:207, figs. 68-72 (female holotype and male paratype from Huangshan,
Anhui, China, in HBI, examined).— Song, Zhu and Chen, 1999:374, figs. 216L—M, 218D, K.
Draconarius aspinatus: Wang, 2002:66.
DIAGNOsIs.— The male of this species is similar to D. altissimus and can only be recognized
by non-toothed conductor. The female is similar to D. wudangensis in having the medially situat-
ed and widely separated epigynal teeth, and similar spermathecal tubes but can be distinguished by
the epigynal teeth position (separated by less than atrial width) (Figs. 13C—E).
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and two
retromarginal teeth. Female epigynal teeth short, situated anterad of atrium; atrium small, slightly
expanded posteriorly; copulatory ducts posteriorly originating, extending mesad of spermathecae;
spermathecal heads slender, situated medially on spermathecae; spermathecal bases small, widely
separated; spermathecal stalks broad, anteriorly expanded and converging (Figs. 13A—B). Male
palp without patellar apophysis; RTA long; lateral tibial apophysis small; cymbial furrow large,
longer than half cymbial length; conductor slender, with large basal lamella; conductor dorsal
apophysis present; embolus posterior in origin; median apophysis spoon-like, elongated (Figs.
13C-—E).
DISTRIBUTION.— China (Anhui) (Map 5).
MATERIAL EXAMINED.— CHINA: Anhui: Huangshan, female holotype and male paratype,
October 24, 1974 (J.F. Wang and C.M. Yin, HBI).
Draconarius baronii (Brignoli, 1978)
Figures 14A—C; Map 5
Coelotes baronii Brignoli, 1978:42, figs. 17-18 (male holotype from Dorjula, Bhutan, in NHMB, examined)
(male only, female paratype is Himalcoelotes brignolii Wang, 2002).
Draconarius baronti: Wang, 2002:66
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Volume 54, No. 26
DIAGNOsIS.— The male of this species is similar to D. tibetensis but can the distinguished by
the long, slender conductor (Figs. 14A—C).
DESCRIPTION.— Described by Brignoli (1978). Chelicerae with three promarginal and two
retromarginal teeth. Male palp with patellar apophysis short; RTA long; lateral tibial apophysis
large, widely separated from RTA; cymbial furrow short; conductor long, slender, with large basal
lamella; conductor dorsal apophysis present; embolus posterior in origin; median apophysis spoon-
like, elongated (Figs. 14A—C). Female unknown.
DISTRIBUTION.— Bhutan (Map 5).
MATERIAL EXAMINED.— BHUTAN: Dorjula: 3100 m, June 6, 1972, male holotype (NHMB,
2302a, 2302b) (female paratype is Himalcoelotes brignolii Wang, 2002).
Draconarius baxiantaiensis Wang, sp. nov.
Figures 1SA—B; 97C; Map 5
Types.— Female holotype and female paratype from Baxiantai, Taibai Mt, Shaanxi, China
(July 13, 1991; X.P. Wang), deposited in IZB.
ETYMOLOGY.— The specific name refers to the type locality.
DIAGNosIs.— The female of this new species is similar to D. potanini in having looped cop-
ulatory ducts and broad, long spermathecae but can be distinguished by the dorsally originating
spermathecal heads and the relatively short copulatory ducts (with only one loop) (Figs. 15SA—B).
FEMALE.— Total length 8.18. Carapace 4.01 long, 2.65 wide. Abdomen 4.17 long, 2.72 wide.
Eye sizes and interdistances: AME 0.07, ALE 0.16, PME 0.12, PLE 0.15; AME-AME 0.1, AME-
ALE 0.08, PME-PME 0.16, PME-PLE 0.19. Leg measurements: I: 8.92 (2.60, 3.22, 1.92, 1.18); II:
8.41 (243, 2.95, 1.88, 1-15); Ml: 7.85 @.22,.2.82,, 1.71, 1-10); TV: 10:8 (2.87, 3-455 22 o7 rea):
Chelicerae with three promarginal and two retromarginal teeth. Female epigynal teeth short, situ-
ated posteriorly near atrium; atrium small, posteriorly situated; copulatory ducts originating poste-
riorly mesad of spermathecae, extending laterally and formed one loop around spermathecae; sper-
mathecal heads long, slender, originating from dorsal side of spermathecae; spermathecal bases
small, widely separated; spermathecal stalks broad, anteriorly extending, slightly converging (Figs.
15A-B).
MALE.— Unknown.
DISTRIBUTION.— China (Shaanxi) (Map 5).
OTHER MATERIAL EXAMINED.— None.
Draconarius bituberculatus (Wang et al., 1990), NEW COMBINATION
Figures 16A—C; Map 6
Coelotes bituberculatus Wang et al., 1990:209, figs. 73-75 (male holotype from Huangshan, Anhui, China, in
HBI, examined).— Song, Zhu and Chen, 1999:374, figs. 218F, M.
DIAGNOsIS.— The male of this species can be easily distinguished from other Draconarius by
the absence of a patellar apophysis, the broad, posteriorly extending conductor (reaching embolic
base), and the retrolaterally extending embolic base (Figs. 16A-C).
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and two
retromarginal teeth. Male palp without patellar apophysis; RTA long, strongly elevated from tibia;
lateral tibial apophysis present; cymbial furrow slightly longer than half cymbium length; conduc-
tor long, broad, posteriorly extending and reaching embolic base, with small basal lamella; conduc-
WANG: EAST ASIA COELOTINE SPIDERS 523
tor dorsal apophysis short; embolus long, retrolateral in origin; median apophysis spoon-like, elon-
gated (Figs. 1}6A—C). Female unknown.
DISTRIBUTION.— China (Anhui) (Map. 6).
MATERIAL EXAMINED: CHINA: Anhui: Huangshan, October 27, 1974, female holotype (C. M.
Yin and J. F. Wang, HBI).
Draconarius brunneus (Hu and Li, 1987), NEW COMBINATION
Map 5
Coelotes brunneus Hu and Li, 1987:277, figs.1—2 (female holotye and female paratype from Yadong, Tibet,
China, in SDU, not examined).— Song, Zhu and Chen, 1999:374, figs. 216V—W;— Hu, 2001:136, figs.
846: 1-2.
DIAGNOSIS — The female of this species is similar to D. disgregus in lacking epigynal teeth and
having broad spermathecae but can be distinguished by the non-convoluted, anteriorly converging
spermathecae.
DESCRIPTION.— See Hu and Li (1987). Chelicerae with three promarginal and two retromar-
ginal teeth. Female without epigynal teeth; spermathecal bases widely separated; spermathecal
stalks broad, anteriorly converging. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 5).
MATERIAL EXAMINED.— None.
Draconarius calcariformis (Wang, 1994)
Figures 17A—D; 97D; Map 6
Coelotes calcariformis Wang, 1994:287, figs. 6-10 (1 male and | female types, no holotype indicated, from
Dabashan, Hubei, China, in HBI, examined).— Song, Zhu and Chen, 1999:374, figs. 217A—B, 218H-I.
Draconarius calcariformis: Wang, 2002:67.
DiAGNosis.— The female is similar to D. colubrinus and D. davidi in having the posteriorly
situated atrium, anteriorly and closely situated epigynal teeth, and similar spermathecal tubes but
can be distinguished by the broad, slightly bifurcate anterior atrial margin and the strongly convo-
luted spermathecae (Figs. 17A—B). The male can be easily identified by the strongly bifurcate con-
ductor and the ventrally and laterally concave tibia (Figs. 17C—D).
DESCRIPTION.— Described by Wang (1994). Chelicerae with three promarginal and two retro-
marginal teeth. Epigynal teeth short, anteriorly situated, adjacent; atrium broad, situated posterior-
ly near epigastric furrow; anterior atrial margin broad, slightly bifurcate; copulatory ducts originat-
ing posteriorly, extending mesad of spermathecae; spermathecal heads slender, situated medially
on spermathecae; spermathecal bases small, widely separated; spermathecal stalks anteriorly
expanded and converging (Figs. 17A—B). Male palp with a short, blunt patellar apophysis; tibia
concave ventrally and laterally; RTA long, almost as long as tibia; lateral tibial apophysis small, sit-
uated anteriorly near RTA; cymbial furrow short; conductor strongly bifurcate, with small basal
lamella; conductor dorsal apophysis present; embolus posterior in origin; median apophysis spoon-
like, small, slightly elongated (Figs. 17C—D).
DISTRIBUTION.— China (Hubei) (Map 6).
MATERIAL EXAMINED.— CHINA: Hubei: Dabashan, Nov. 10, 1990, | male and 1 female types
(J.F. Wang, HBI).
524 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconarius capitulatus Wang, sp. nov.
Figures 18A—B; Map 6
Types.— Female holotype and 5 female paratypes from Pianma Yakou, pass over
Gaoligongshan, at elev. 3200m, 25°58’N, 98°41’E, Nujiang Prefecture, Yunnan, China (October 11,
1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (male holotype and 3 female
paratypes) and CAS (2 female paratypes); 1 female paratype from Nujiang Prefecture, Nujiang
State Nature Reserve, No. 12 Bridge Camp area, 16.3 air km W of Gongshan, N27.715°/E98.502°,
2775m, Gaoligong Shan, Yunnan, China (July 15-19, 2000; H.M. Yan, D. Kavanaugh, C.E.
Griswold, H.B. Liang, D. Ubick, and D.Z. Dong), deposited in CAS.
ETYMOLOGY.— The specific name refers to the prominent spermathecal heads.
DIAGNOsIs.— The female of this species is similar to D. pseudocapitulatus in lacking epigy-
nal teeth and having short, medially lobed spermathecae but can be distinguished by the closely sit-
uated, not anteriorly expanded spermathecae (Figs. 18A—B).
FEMALE.— Total length 10.6. Carapace 5.20 long, 3.60 wide. Abdomen 5.40 long, 3.60 wide.
Eye sizes and interdistances: AME 0.16, ALE 0.27, PME 0.25, PLE 0.25; AME-AME 0.12, AME-
ALE 0.10, PME-PME 0.15, PME-PLE 0.28, AME-PME 0.20. Leg measurements: I: 13.1 (3.80,
4.70, 3.04, 1.60); Il: 12.3 (3.60, 4.42, 2.80, 1.50); HI: 10.3 (2.80, 3.60, 2.60, 1.30); IV: 14.3 (4.00,
4.80, 3.80, 1.70). Chelicerae with three promarginal and two retromarginal teeth. Female epigynal
teeth absent; atrium posteriorly situated; copulatory ducts originating posteriorly, extending mesad
of spermathecae, slightly separated; spermathecal heads large, situated anteriorly on inner side of
spermathecae; spermathecal bases small, widely separated; spermathecal stalks short, broad (Figs.
18A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 6).
OTHER MATERIAL EXAMINED.— None.
Draconarius carinatus (Wang et al., 1990), NEW COMBINATION
Figures 19A—E; Map 7
Coelotes carinatus Wang et al., 1990:211, figs. 76-80 (female holotype, male and female paratypes from
Huangshan, Anhui, China, in HBI, examined).— Song, Zhu and Chen, 1999:374, figs. 217C—D, 219A—H.
DIAGNOsISs.— This species can be easily distinguished from other coelotines by the broad, lat-
erally originating, medially extending copulatory ducts of female (Figs. 19A—B), and by the
absence of patellar apophysis, the strongly bifurcate median apophysis, and the broad embolus of
male (Figs. 19C—E).
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and two
retromarginal teeth. Female epigynal teeth large, situated posteriorly on anterior atrial margin; atri-
um small, posteriorly situated; copulatory ducts originating laterally, looped around spermathecae
and extending medially; spermathecal heads small, anteriorly situated; spermathecae small, wide-
ly separated (Figs. 19A—B). Male palp lacking patellar apophysis; RTA strongly elevated from
tibia; lateral tibial apophysis widely separated from RTA; cymbial furrow short; conductor broad,
deeply grooved, with small basal lamella; conductor dorsal apophysis present; embolus broad, pos-
terior in origin; median apophysis strongly bifurcate, not spoon-like (Figs. 19C—E).
DISTRIBUTION.— China (Anhui) (Map 7).
MATERIAL EXAMINED.— CHINA: Anhui: Huangshan, October 27, 1974, female holotype, 1
male and | female paratypes (C.M. Yin and J.F. Wang, HBI).
WANG: EAST ASIA COELOTINE SPIDERS 525
Draconarius chaigiaoensis (Zhang, Peng and Kim, 1997), NEw COMBINATION
Figures 20A—C; 97E; Map 6
Coelotes chaigiaoensis Zhang, Peng and Kim, 1997:291, figs. 1-3 (male holotype from Chaigiao, Ningbo,
Zhejiang, China, in HBI, examined).
DIAGNOSIS.— The male of this species can be easily distinguished from other coelotines by
the absence of a patellar apophysis, the short cymbial furrow, and the presence of short, broad con-
ductor (Figs. 20A-C).
DESCRIPTION.— Described by Zhang, Peng and Kim (1997). Chelicerae with three promargin-
al and two retromarginal teeth. Male palp without patellar apophysis; RTA short, approximately
half tibial length; lateral tibial apophysis widely separated from RTA; cymbial furrow short; con-
ductor short, broad, with large dorsal edge and reduced basal lamella; conductor dorsal apophysis
present; embolus prolateral in origin; median apophysis spoon-like, slightly elongated (Figs.
20A—C). Female unknown.
DISTRIBUTION.— China (Zhejiang) (Map 6).
MATERIAL EXAMINED.— CHINA: Zhejiang: Ningbo, Chaiqiao, November 20, 1991, female
holotype (Y.J. Zhang, HBI).
Draconarius cheni (Platnick, 1989)
Figures 21A—B; Map 7
Coelotes saxatilis Chen, 1984:2, figs. 5-6 (female holotype and female paratype from Huanglongdong,
Hangzhou, Zhejiang, China, in HTC, examined).— Chen and Zhang, 1991:189, figs. 188.1—2. (specific
name preoccupied by Balckwall, 1833).
Coelotes cheni Platnick, 1989:422 (replacement name).— Song, Zhu and Chen, 1999:374, figs. 217E-F.
Draconarius cheni: Wang, 2002:67.
DIAGNOSIS.— The female of this species is similar to D. arcuatus in having a broad, medially
situated copulatory ducts but can be distinguished by the small atrium (same width as epigynal
teeth distance) and the indistinct spermathecal heads (Figs. 21A—B).
DESCRIPTION.— The female was described by Chen (1984). Chelicerae with three promargin-
al and two retromarginal teeth. Female epigynal teeth short, situated anterad of atrium, separated
by atrial width; atrium small; copulatory ducts broad, originating posteriorly, extending anteriorly
mesad of spermathecae, slightly folded; spermathecal heads not visible from dorsal view; sper-
mathecal bases widely separated; spermathecal stalks long, slender, extending laterally and then
slightly converging anteriorly (Figs. 21A—B). Male unknown.
DISTRIBUTION.— China (Zhejiang) (Map 7).
MATERIAL EXAMINED.— CHINA: Zhejiang: Hangzhou, March 1977, female holotype and 1
female paratype (Z.F. Chen, HTC).
Draconarius colubrinus Zhang, Zhu and Song, 2002
Map 7
Draconarius colubrinus Zhang, Zhu and Song, 2002:52, figs. 1-4 (male holotype and three female paratypes
from Muyu, Shennongjia, Hubei, China, in HU, not examined).
DIAGNOsIs.— The female is similar to D. calcariformis in having the posteriorly situated atri-
um, anteriorly and closely situated epigynal teeth, and similar spermathecal tubes, but can be dis-
526 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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tinguished by the broad, moderately separated spermathecal bases. The male is similar to D. neix-
iangensis but can be recognized by the slender embolic apex.
DESCRIPTION.— See Zhang, Zhu and Song (2002). Cheliceral promargin with three teeth,
retromargin with two. Epigynal teeth short, anteriorly situated, close together; atrium small, poste-
riorly situated, near epigastric furrow; copulatory ducts originating posteriorly, extending mesad of
spermathecae; spermathecal heads situated laterally on spermathecae; spermathecal bases small,
widely separated; spermathecal stalks broad, anteriorly expanded and converging. Male palp with-
out patellar apophysis; RTA long; lateral tibial apophysis large; cymbial furrow short; conductor
broad, with basal lamella small; conductor dorsal apophysis small; embolus broad, posterior in ori-
gin, with slender apex; median apophysis spoon-like, elongated.
DISTRIBUTION.— China (Hubei) (Map. 7).
MATERIAL EXAMINED.— None.
Draconarius coreanus (Paik and Yaginuma, 1969)
Figures 22A—E; Map 7
Coelotes coreanus Paik and Yaginuma, 1969:837, figs. 62-64 (types deposited in the National Science
Museum, Tokyo, not examined).— Paik, 1978:337, figs. 149.1-3.
Draconarius coreanus: Wang, 2002:67.
DIAGNOosIs.— The female is similar to D. wudangensis in having the medially situated and
widely separated epigynal teeth, and similar spermathecal tubes but can be distinguished by the
broad atrium and the slender spermathecal bases (Figs. 22A—B). The male is similar to D. wudan-
gensis but can be recognized by the long RTA (more than half tibial length) and the broad median
apophysis (Figs. 22C-E).
DESCRIPTION.— See Paik, Yaginuma and Namkung (1969). Cheliceral promargin with three
teeth, retromargin with two. Epigynal teeth short, widely separated; atrium small, situated posteri-
orly near epigastric furrow; anterior atrial margin broad, membranous; copulatory ducts originat-
ing posteriorly near epigastric furrow, extending mesad of spermathecae; spermathecal heads situ-
ated medially on spermathecae; spermathecal bases small; spermathecal stalks broad, anteriorly
expanded and converging (Figs. 22A—B). Male palp with patellar apophysis long; RTA long; later-
al tibial apophysis present; cymbial furrow more than half cymbial length; conductor short, slen-
der, with large basal lamella; conductor dorsal apophysis slender; embolus posterior in origin;
median apophysis spoon-like, slightly elongated (Figs. 22C—E).
DISTRIBUTION.— South Korea (Map. 7).
MATERIAL EXAMINED.— SOUTH KOREA: Mt. Kwan-ak, May 5, 1991, 1 female (Cheol-hoe
Jung, KAT); Mt. Hallason, August 7, 1984, 1 male (J.P. Kim, KAI).
Draconarius curiosus Wang, sp. nov.
Figures 23A—D; Map 7
Types.— Male holotype and female paratype from Luoshuidong, 28 air km E TengChong,
24°57’N, 98°45’E, native forest, 2300m, Baoshan Prefecture, Yunnan, China (October 26-31,
1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI; 1 female paratype from Pass
over Gaoligongshan at 2100 m, Nankang, 36 air km SE TengChong, 24°50’N, 98°47’E, native for-
est, Baoshan Prefecture, Yunnan, China (November 4—7, 1998: C. Griswold, D. Kavanaugh, C-L.
Long), deposited in HBI; 2 male and 15 female paratypes from Luoshuidong, Baoshan, Yunnan,
WANG: EAST ASIA COELOTINE SPIDERS 527
China (October 26-31, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (1 male
and 7 females) and CAS (1 male and 8 females).
EtyMoLoGy.— The specific name refers to its odd palpal structure.
DIAGNosIs.— The female of this new species is similar to D. griswoldi by the absence of epig-
ynal teeth and the laterally situated copulatory ducts but can be distinguished by the looped sper-
mathecae (Fig. 23B). The male of this species can be easily recognized from all other coelotines
by the absence of a median apophysis, the broad embolus, and strongly expanded embolic apex
(Figs. 23C, D).
FEMALE.— Total length 6.80. Carapace 3.20 long, 2.14 wide. Abdomen 3.60 long, 2.50 wide.
Eye sizes and interdistances: AME 0.09, ALE 0.19, PME 0.19, PLE 0.19; AME-AME 0.09, AME-
ALE 0.04, PME-PME 0.08, PME-PLE 0.12, AME-PME 0.11. Leg measurements: I: 7.06 (2.12,
2.46, 1.60, 0.88); Il: 6.44 (1.92, 2.18, 1.40, 0.94); III: 6.06 (1.64, 2.00, 1.52, 0.90); IV: 8.48 (2.20,
2.80, 2.34, 1.14). Chelicerae with three promarginal and two retromarginal teeth. Epigynal teeth
absent; atrium broad; copulatory ducts broad, originating posteriorly, extending laterad of sper-
mathecae, anteriorly converging; spermathecal heads situated inside spermathecal loops; spermath-
ecal bases large, widely separated; spermathecal stalks extending with one loop (Figs. 23A—B).
MALE.— Total length 5.62. Carapace 2.90 long, 2.00 wide. Abdomen 2.72 long, 1.60 wide.
Eye sizes and interdistances: AME 0.09, ALE 0.16, PME 0.18, PLE 0.18; AME-AME 0.05, AME-
ALE 0.03, PME-PME 0.06, PME-PLE 0.07, AME-PME 0.09. Leg measurements: I: 7.82 (2.22,
2.74, 1.80, 1.06); II: 7.10 (2.04, 2.40, 1.66, 1.00); III: 6.46 (1.82, 2.02, 1.72, 0.90); IV: 8.72 (2.34,
2.68, 2.52, 1.18). Chelicerae with three promarginal and two retromarginal teeth. Male palp with
patellar apophysis present; RTA long; lateral tibial apophysis small; cymbial furrow short; conduc-
tor short, deeply grooved, with basal lamella small; conductor dorsal apophysis small; embolus
posterior in origin, broad, with strongly expanded apex; median apophysis absent (Figs. 23C, D).
DISTRIBUTION.— China (Yunnan) (Map 7).
OTHER MATERIAL EXAMINED.— None.
Draconarius davidi (Schenkel, 1963)
Figures 24A— B; Map 8
Coelotes davidi Schenkel, 1963:283, fig. 159 (female holotype from Inkiaphou, Shensi, China, in MNHN,
examined).— Song, Zhu and Chen, 1999:374.
Draconarius davidi: Wang, 2002:67.
DIAGNOsIs.— The female is similar to D. calcariformis in having the posteriorly situated atri-
um, anteriorly and closely situated epigynal teeth, and similar spermathecal tubes but can be dis-
tinguished by the broad spermathecal bases (Figs. 24A—B).
DESCRIPTION.— See Schenkel (1963). Chelicerae with three promarginal and two retromar-
ginal teeth. Female epigynal teeth short, anteriorly situated, close together; atrium small, posteri-
orly situated, near epigastric furrow; copulatory ducts originating posteriorly, extending mesad of
spermathecae; spermathecal heads not visible on the examined specimen; spermathecal bases
small, widely separated; spermathecal stalks broad, anteriorly expanded and converging (Figs.
24A-B). Male unknown.
DISTRIBUTION.— China (Shaanxi) (Map 8).
MATERIAL EXAMINED.— CHINA: Shaanxi (Shensi): Inkiaphou, | female type, collected in
1873 (A. David, MNHN, B2011 bis.).
528 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconarius denisi (Schenkel, 1963) NEW COMBINATION
Figures 25A—B; Map 8
Coelotes denisi Schenkel, 1963:285, fig. 160 (female holotype from Lo Thoei-Tong, Yunnan, China, in
MNHBN, examined).
DIAGNosIs.— The female of this species is similar to D. adligansus but can be separated by
the anteriorly situated, widely separated epigynal teeth (away from atrium, separated by at least
atrial width) and the small spermathecae (Figs. 25A—B).
DESCRIPTION.— See Schenkel (1963). Chelicerae with three promarginal and two retromar-
ginal teeth. Female epigynal teeth short, situated anteriorly, widely separated; atrium small; copu-
latory ducts large, anteriorly extending, close together; spermathecal heads situated laterally; sper-
mathecae short, widely separated (Figs. 25A—B). Male unknown.
DISTRIBUTION.— China (Yunnan) (Map 8).
MATERIAL EXAMINED.— CHINA: Lo Thoei Tong (Yunnan?), March 2, 1925, female holotype
(MNHN, B2011 bis).
Draconarius digitusiformis (Wang et al., 1990), NEw COMBINATION
Figures 26A—E; 97F; Map 8
Coelotes digitusiformis Wang et al., 1990:205, figs. 63-67 (1 male and 2 female types, holotype not indicat-
ed, from Zhong Village, Ling County, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:
374, figs. 217K—-L, 219F, M.
Coelotes shuangpaiensis Peng, Gong and Kim, 1996:20, figs. 15-18 (male holotype from Shuangpai, Hunan,
China, in HBI, examined).— Song, Zhu and Chen, 1999:378, figs. 226U, 228C. NEw SYNONYMY.
DIAGNOsIs.— This species can be easily distinguished by the closely situated epigynal teeth,
the anteriorly situated, strongly convoluted copulatory ducts of female (Figs. 26A—B), and by the
broad, anteriorly extending, slightly spiraled conductor of male (Figs. 26C—E).
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and two
retromarginal teeth. Female epigynal teeth situated anteriorly, close together; atria small, widely
separated; copulatory ducts broad, strongly convoluted, extending anterad of spermathecae; sper-
mathecal heads situated posterad of copulatory ducts; spermathecal bases broad, close together;
spermathecal stalks short, broad, extending laterally (Figs. 204—B). Male palp with patellar apoph-
ysis long, broad; RTA almost as long as tibia; lateral tibial apophysis small; cymbial furrow more
than half cymbial length; conductor short, broad, form broad groove and slightly spiraled apex,
with large basal lamella; conductor dorsal apophysis small; embolus posterior in origin; median
apophysis small, spoon-like (Figs. 26C—E).
DISTRIBUTION.— China (Hunan) (Map 8).
MATERIAL EXAMINED.— CHINA: Hunan: Ling County, Zhong Village, December 15, 1982, 1
male and 2 female types (J.F. Wang, HBI); Shuangpai, October 3, 1993, male holotype of Coelotes
shuangpaiensis (C.L. He, HBI).
Draconarius disgregus Wang, sp. nov.
Figures 27A—D; Map 8
Typrs.— Male holotype and female paratype from native forest in Gaoligongshan at 9.5 road
km ESE Pianma, 25°59’N, 98°40’E, el. 2500m, Nujiang Prefecture, Yunnan, China (October
15-18, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI; 2 male and 5 female
WANG: EAST ASIA COELOTINE SPIDERS 529
paratypes from native forest in Gaoligongshan at 9.5 road km ESE Pianma, 25°59’/N, 98°40’E, el.
2500m, Nujiang Prefecture, Yunnan, China (October 15-18, 1998; C. Griswold, D. Kavanaugh, C-
L. Long), deposited in HBI (1 male and 2 females) and CAS (1 male and 3 females).
ETYMOLOGY.— The specific name refers to its difference from other Draconarius in both
male and female genitalia.
DIAGNosIs.— The female of this species is similar to D. brunneus in lacking epigynal teeth
and having broad spermathecae but can be distinguished by the convoluted, anteriorly diverging
spermathecae. (Figs. 27A—B). The males can be easily distinguished from other coelotines by the
short, slightly curved patellar apophysis and the broad, not spoon-like median apophysis (Figs.
27C-D).
FEMALE.— Total length 11.4. Carapace 5.80 long, 3.70 wide. Abdomen 5.60 long, 4.00 wide.
Eye sizes and interdistances: AME 0.17, ALE 0.26, PME 0.24, PLE 0.25; AME-AME 0.15, AME-
ALE 0.12, PME-PME 0.15, PME-PLE 0.25, AME-PME 0.20. Leg measurements: I: 13.8 (4.00,
4.80, 3.10, 1.90); II: 12.4 (3.60, 4.20, 2.80, 1.80); II: 10.9 (3.00, 3.60, 2.80, 1.50); IV: 14.7 (4.00,
4.80, 4.00, 1.90). Chelicerae with three promarginal and two retromarginal teeth. Epigynum lack-
ing epigynal teeth; atrium small; copulatory ducts short, invisible from dorsal view; spermathecal
heads large, anteriorly situated; spermathecal bases widely separated; spermathecal stalks broad,
anteriorly extending and slightly converging (Figs. 27A—B).
MALE.— Total length 9.90. Carapace 5.10 long, 3.80 wide. Abdomen 4.80 long, 3.40 wide.
Eye sizes and interdistances: AME 0.17, ALE 0.24, PME 0.21, PLE 0.23; AME-AME 0.10, AME-
ALE 0.05, PME-PME 0.11, PME-PLE 0.19, AME-PME 0.20. Leg measurements: I: 17.3 (4.60,
5.90, 4.40, 2.40); I: 15.7 (4.20, 5.10, 4.10, 2.30); HI: 13.4 (3.60, 4.20, 3.70, 1.90); IV: 17.1 (4.60,
5.20, 5.00, 2.30). Chelicerae with three promarginal and two retromarginal teeth. Male palp with
patellar apophysis short, slightly curved dorsally; RTA almost as long as tibia; lateral tibial apoph-
ysis small; cymbial furrow short; conductor with broad apex and small basal lamella; conductor
dorsal apophysis small; embolus prolateral in origin; median apophysis broad, membranous, not
spoon-like (Figs. 27C—D).
DISTRIBUTION.— China (Yunnan) (Map 8).
OTHER MATERIAL EXAMINED.— None.
Draconarius dissitus Wang, sp. nov.
Figures 28A—B; Map 9
TyprEs.— Female holotype, 3 female paratypes from Yupin area (S. Tibet), Tibet, China (May,
1998: G Schaller), deposited in AMNH.
ETYMOLOGY. — The specific name refers to the widely separated spermathecae.
DIAGNOsIs.— The female of this species is similar to D. capitulatus and D. pseudocapitulatus
but can the distinguished by the absence of medially expanded spermathecal lobes and the pres-
ence of small, widely separated spermathecae (Figs. 283A—B).
FEMALE.— Total length 13.8. Carapace 5.80 long, 4.00 wide. Abdomen 8.00 long, 5.20 wide.
Eye sizes and interdistances: AME 0.18, ALE 0.26, PME 0.22, PLE 0.24; AME-AME 0.12, AME-
ALE 0.10, PME-PME 0.22, PME-PLE 0.25, AME-PME 0.20. Leg measurements: I: 13.0 (3.80,
4.64, 2.88, 1.68); II: 12.1 (3.44, 4.16, 2.88, 1.60); III: 11.0 (3.20, 3.60, 2.80, 1.36); IV: 14.2 (3.84,
4.72, 3.88, 1.80). Chelicerae with three promarginal, and two retromarginal teeth. Epigynal teeth
absent; atrium small; epigynal hoods distinct, situated anteriorly; copulatory ducts short, situated
mesad of spermathecae; spermathecal heads small, situated mesad of spermathecae; spermathecae
small, slightly extending anteriorly, widely separated (Figs. 23A—B).
530 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
MALE.— Unknown.
DISTRIBUTION.— China (Tibet) (Map 9).
OTHER MATERIAL EXAMINED.— None.
Draconarius dubius Wang, sp. nov.
Figures 29A—D; Map 9
TypEs.— Male holotype and female paratype from Pianma Yakou, pass over Gaoligongshan,
at elev. 3200m, 25°58’N, 98°41’E, Nujiang Prefecture, Yunnan, China (October 11, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI; 7 female paratypes from Pianma Yakou,
Nujiang, Yunnan, China (October 11, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in
HBI (3 females) and CAS (4 females); 4 male paratypes from native forest in Gaoligongshan at 9.5
road km ESE Pianma, 25°59’N, 98°40’E, el. 2500m, Nujiang Prefecture, Yunnan, China, (October
15-18, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (2 males) and CAS (2
males).
ETYMOLOGY.— The specific name refers to the unusual morphology compare to other
Draconarius: the not spoon-like median apophysis and the absence of epigynal teeth.
DIAGNosis.— The female of this species is similar to D. patellabifidus and D. simplicidens by
lacking epigynal teeth and having broad spermathecae but can be recognized from D. simplicidens
by the anteriorly situated spermathecal heads, from D. patellabifidus by the broader spermathecal
bases and the anteriorly diverging spermathecae (Figs. 29A—B). Male can be recognized by the not
spoon-like median apophysis, the bifurcated conductor, and the long patellar apophysis (longer
than patellar length) (Figs. 29C—D).
FEMALE.— Total length 8.00. Carapace 4.00 long, 2.90 wide. Abdomen 4.00 long, 2.70 wide.
Eye sizes and interdistances: AME 0.15, ALE 0.21, PME 0.19, PLE 0.20; AME-AME 0.10, AME-
ALE 0.05, PME-PME 0.13, PME-PLE 0.18, AME-PME 0.20. Leg measurements: I: 10.3 (2.90,
3.70, 2.30, 1.40); I: 9.60 (2.70, 3.30, 2.20, 1.40); III: 9.00 (2.40, 3.00, 2.30, 1.30); IV: 12.0 (3.20,
3.80, 3.30, 1.70). Chelicerae with three promarginal, and two retromarginal teeth. Epigynal teeth
absent; atrium large; copulatory ducts small, situated mesad of spermathecae; spermathecal heads
large, situated anteriorly mesad of spermathecae; spermathecal bases broader than stalks; sper-
mathecal stalks anteriorly diverging (Figs. 29A—B).
MALE.— Total length 6.80. Carapace 3.60 long, 2.66 wide. Abdomen 3.20 long, 3.00 wide.
Eye sizes and interdistances: AME 0.11, ALE 0.20, PME 0.18, PLE 0.19; AME-AME 0.05, AME-
ALE 0.05, PME-PME 0.08, PME-PLE 0.14, AME-PME 0.13. Leg measurements: I: 11.3 (3.10,
3.94, 2.60, 1.70); II: 10.2 (2.80, 3.40, 2.46, 1.52); Il: 9.40 (2.70, 2.80, 2.50, 1.40); IV: 12.1 (3.20,
3.80, 3.50, 1.60). Chelicerae with three promarginal, and two retromarginal teeth. Male palp with
patellar apophysis longer than patellar length; RTA long; lateral tibial apophysis large, widely sep-
arated from RTA; cymbial furrow short; conductor broad, bificated, with small basal lamella; con-
ductor dorsal apophysis present; embolus prolateral in origin; median apophysis simple, not spoon-
like (Figs. 29C—D).
DISTRIBUTION.— China (Yunnan) (Map 9).
OTHER MATERIAL EXAMINED.— None.
Draconarius episomos Wang, sp. nov.
Figures 30A—B; Map 10
TypEs.— Female holotype and female paratype from Pianma Yakou, pass over Gaoligong-
WANG: EAST ASIA COELOTINE SPIDERS 531
shan, at elev. 3200m, 25°58’N, 98°41’E, Nujiang Prefecture, Yunnan, China (October 11, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (female holotype) and CAS (female
paratype).
EtyMOLOoGyY.— The specific name refers to the large spermathecae.
DiaGNosts.— The female is similar to D. himalayaensis, D. altissimus and D. subtitanus in
having the short epigynal teeth and rounded, closely situated spermathecae but can be distinguished
by the widely separated epigynal teeth (at least one and half atrial width) and the medially situat-
ed spermathecal heads. (Figs. 30A—B).
FEMALE.— Total length 10.7. Carapace 4.68 long, 2.99 wide. Abdomen 5.98 long, 3.64 wide.
Eye sizes and interdistances: AME 0.11, ALE 0.16, PME 0.14, PLE 0.16; AME-AME 0.11, AME-
ALE 0.13, PME-PME 0.20, PME-PLE 0.22, AME-PME 0.13. Leg measurements: I: 11.0 (3.23,
pote. fo): 1 919312786) 3:38; 2.217) 1-48): Ml: OA 2:5523.09)2:345 1:43); IV-12 562283
4.16, 3.38, 1.66). Chelicerae with three promarginal, and two retromarginal teeth. Epigynal teeth
short, widely separated; atrium small, near epigastric furrow; copulatory ducts small, situated
mesad of spermathecae; spermathecal heads small, situated anteriorly mesad of spermathecae;
spermathecal bases small, widely separated; spermathecal stalks broad, anteriorly expanded and
converging (Figs. 30A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 10).
OTHER MATERIAL EXAMINED.— None.
Draconarius everesti (Hu, 2001), NEW COMBINATION
Map 9
Coelotes everesti Hu, 2001:145, figs.8-55:1—3 (male holotye from Ang-Ren, Tibet, China, in SDU, not exam-
ined).
DIAGNOSIS — The male of this species is similar to D. singulatus in having a short cymbial fur-
row and prolaterally originating embolus but can be distinguished by the short RTA.
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Male palp with patellar apophysis present; RTA short, much less than half tibial length; cym-
bial furrow short; conductor short; embolus prolateral in origin; median apophysis spoon-like,
slightly elongated. Female unknown.
DISTRIBUTION.— China (Tibet) (Map 9).
MATERIAL EXAMINED.— None.
Draconarius griswoldi Wang, sp. nov.
Figures 31A—D; Map 11
TyprEs.— Female holotype and 2 female paratypes from Pianma Yakou, pass over Gaoligong-
shan, at elev. 3200m, 25°58’N, 98°41’E, Nujiang Prefecture, Yunnan, China (October 11, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (holotype female) and CAS (paratype
females); 2 male paratypes from native forest in Gaoligongshan at 9.5 road km ESE Pianma,
25°59’N, 98°40’E, el. 2500m, Nujiang Prefecture, Yunnan, China (October 15—18; C. Griswold, D.
Kavanaugh, C-L. Long), deposited in HBI (1 male) and CAS (1 male).
ETYMOLOGY.— The specific name is a patronym in honor of Dr. Charles E. Griswold, the col-
lector of the specimens.
532 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DIAGNOsIS.— The female of this new species is similar to D. curiosus in having the laterally
extending copulatory ducts and lacking epigynal teeth but can be easily distinguished by the ante-
riorly situated spermathecal heads and the unlooped spermathecae (Figs. 31A—B). The male is sim-
ilar to D. agrestis in having a small patellar apophysis, an elongated cymbial furrow, and a poste-
riorly originated, long embolus but can be distinguished by the slender embolus, the sharp conduc-
tor apex, and the slightly spoon-like median apophysis (Figs. 31C—D).
FEMALE.— Total length 5.60. Carapace 2.60 long, 1.80 wide. Abdomen 3.00 long, 2.00 wide.
Eye sizes and interdistances: AME 0.08, ALE 0.18, PME 0.15, PLE 0.18; AME-AME 0.03, AME-
ALE 0.02, PME-PME 0.04, PME-PLE 0.10, AME-PME 0.10. Leg measurements: I: 5.90 (1.72,
2.14, 1.24, 0.80); II: 5.40 (1.58, 1.90, 1.20, 0.72); Ill: 5.10 (1.40, 1.70, 1.30, 0.70); IV: 6.86 (1.80,
2.30, 1.86, 0.90). Chelicerae with three promarginal, and two retromarginal teeth. Epigynal teeth
absent; atrium large; copulatory ducts originating posteriorly laterad of spermathecae, connected to
spermathecae anteriorly; spermathecal heads small, situated anteriorly; spermathecal bases broad,
widely separated; spermathecal stalks long, anteriorly converging (Figs. 31A—B).
MALE.— Total length 5.36. Carapace 2.76 long, 1.80 wide. Abdomen 2.60 long, 1.60 wide.
Eye sizes and interdistances: AME 0.07, ALE 0.17, PME 0.15, PLE 0.15; AME-AME 0.04, AME-
ALE 0.03, PME-PME 0.05, PME-PLE 0.10, AME-PME 0.07. Leg measurements: I: 7.26 (2.16,
2.50, 1.60, 1.00); Il: 6.60 (1.92, 2.20, 1.50, 0.98); III: 5.54 (1.64, 1.60, 1.50, 0.80); IV: 8.10 (2.20,
2.60, 2.20, 1.10). Chelicerae with three promarginal, and two retromarginal teeth. Male palp with
patellar apophysis small; RTA long; lateral tibial apophysis present; cymbial furrow more then half
cymbial length; conductor broad, with slender apex and large basal lamella; conductor dorsal
apophysis broad; embolus posterior in origin; median apophysis broad, elongated, slightly spoon-
like (Figs. 31C—D).
DISTRIBUTION.— China (Yunnan) (Map 11).
OTHER MATERIAL EXAMINED.— None.
Draconarius funitushanensis (Hu, Wang and Wang, 1991), NEw COMBINATION
Map 9
Coelotes funiushanensis Hu, Wang and Wang, 1991:41, figs. 14-17 (female holotype, male and female
paratypes from Yaochanggou, Neixiang, Henan, China, in SDU, not examined).— Song, Zhu and Chen,
1999:375, figs. 217Q-R, 222B, 223G
DIAGNOsIS.— The female is similar to D. hui but can be distinguished by the closely situated
spermathecal bases. The male can be easily recognized by the long, lobed patellar apophysis.
DESCRIPTION.— See Hu, Wang and Wang (1991). Chelicerae with three promarginal, and two
retromarginal teeth. Female epigynal teeth near atrium; atrium small; spermathecal bases small,
close together; spermathecal stalks broad, anteriorly expanded, close together. Male palpal patellar
apophysis large, strongly lobed; RTA long; embolus posterior in origin.
DISTRIBUTION.— China (Henan) (Map 9).
MATERIAL EXAMINED.— None.
Draconarius gurkha (Brignoli, 1976)
Figures 32A—B; Map 10
Coelotes gurkha Brignoli, 1976:239, figs. 13-14 (female holotype from Yak-Alm, Taboche, Nepal, in IZI,
Np61-—77, examined).
Draconarius gurkha: Wang, 2002:67.
WANG: EAST ASIA COELOTINE SPIDERS 33
DIAGNosIs.— The female is similar to D. singulatus by the indistinct spermathecal heads but
can be distinguished by the small atrium and the indistinct copulatory ducts (Figs. 32A—B).
DESCRIPTION.— See Brignoli (1976). Chelicerae with three promarginal and two retromargin-
al teeth. Female epigynal teeth situated near atrium; atrium small, near epigastric furrow; copula-
tory ducts invisible from dorsal view; spermathecal bases narrow, widely separated; spermathecal
stalks broad, rounded (Figs. 32A—B). Male unknown.
DISTRIBUTION.— Nepal (Fig. Map 10).
MATERIAL EXAMINED.— NEPAL: Taboche (= Taweche), Yak-Alm, Zwergstranchheideboden,
4550 m, May 31, 1961, female holotype (H. Janetschek, IZI, Np61—77); Mingbo-Tal beim Airstrip
der Hillary-Makalu Expedition, 4800 m, May 28, 1961, female type of Coelotes lama Brignoli,
1976 (IZI, Np61-—71).
Draconarius gyriniformis (Wang and Zhu, 1991), NEW COMBINATION
Map 10
Coelotes gyriniformis Wang and Zhu, 1991:4, figs. 11-12 (female holotype, 1 female paratype from
Kangding, Sichuan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:375, figs. 217Y-—Z.
DiAGNosIs.— The female of this species is similar to D. wudangensis in having medially sit-
uated and widely separated epigynal teeth, and similar spermathecal tubes but can be distinguished
by laterally situated spermathecal heads.
DESCRIPTION.— See Wang and Zhu (1991). Chelicerae with three promarginal, and two retro-
marginal teeth. Female epigynal teeth widely separated, anterior of atrium; atrium small, near epi-
gastric furrow; spermathecal bases small, widely separated; spermathecal stalks broad, anteriorly
expanded and converging. Male unknown.
DISTRIBUTION.— China (Sichuan) (Map 10).
MATERIAL EXAMINED.— None.
Draconarius haopingensis Wang, sp. nov.
Figures 33A—B; 96A; Map 10
TypEs.— Female holotype and female paratype from Haoping, Taibaishan, Shaanxi, China
(August 11, 1989; X.P. Wang), deposited in IZB.
ETYMOLOGY.— The specific name refers to the type locality.
DIAGNOsIs.— The female of this new species is similar to D. molluscus by lacking epigynal
teeth and having laterally situated epigynal hoods but can be distinguished by the single loop of
copulatory ducts and the smooth, non-convoluted spermathecal distal ends (Figs. 33A—B).
FEMALE.— Total length 5.19. Carapace 2.28 long, 1.76 wide. Abdomen 2.91 long, 1.89 wide.
Eye sizes and interdistances: AME 0.12, ALE 0.14, PME 0.14, PLE 0.15; AME-AME 0.08, AME-
ALE 0.05, PME-PME 0.11, PME-PLE 0.13. Leg measurements: I: 7.18 (1.94, 2.61, 1.69, 0.94); IT:
6-43 (1-381, 2:31, 1:40, 0.79); Ill: 5.68.64, 1.85, 1.40; 0:79); IV: 7.76 (2.20; 2:59; 2:06, 0:91).
Chelicerae with three promarginal and three retromarginal teeth. Epigynal teeth absent; atrium
large; epigynal hoods situated posteriorly, laterad of atrium; copulatory ducts broad, originating
posteriorly, extending mesad of spermathecae, looped around spermathecae; spermathecal bases
widely separated; spermathecal stalks long, anteriorly converging (Figs. 33A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Shaanxi) (Map 10).
OTHER MATERIAL EXAMINED.— None.
534 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconarius hangzhouensis (Chen, 1984), NEw COMBINATION
Figures 34A—B; Map 11
Coelotes hangzhouensis Chen, 1984:1, figs.1-2 (female holotype from Yunqi, | female paratype from
Huanglongdong, Hangzhou, Zhejiang, China, in HTC, examined).— Chen and Zhang, 1991:188, figs.
186.1-2;— Song, Zhu and Chen, 1999:375, figs. 217A—B.
DIAGNOSIS.— This species can be easily recognized by the trifurcated, less sclerotized anteri-
or atrial margin, the broad, anteriorly extending copulatory ducts, and the long, large spermathecal
heads of the female (Fig. 34A—B).
DESCRIPTION.— See Chen (1984). Chelicerae with three promarginal and two retromarginal
teeth. Female epigynal teeth short, situated anteriorly; anterior atrial margin modified into three
broad, less sclerotized pieces; atrium small, near epigastric furrow; copulatory ducts broad, origi-
nating posteriorly, mesad of spermathecae, extending anteriorly anterad of spermathecae; sper-
mathecal heads large, anteriorly extending; spermathecae broad, widely separated (Figs. 34A—B).
Male unknown.
DISTRIBUTION.— China (Zhejiang) (Map 11).
MATERIAL EXAMINED.— CHINA: Zhejiang: Hangzhou, Yungi, October 7, 1981, female holo-
type (Z.F. Chen, HTC); Huanglongdong, October 7, 1979, | female paratype (Z.F. Chen, HTC).
Draconarius himalayaensis (Hu, 2001), NEW COMBINATION
Map 11
Coelotes himalayaensis Hu, 2001:134, figs. 8-45:1—4 (female holotype, male and female paratypes from
Linzhi, Tibet, China, in SDU, not examined).
DiAGnosis.— The female is similar to D. altissimus and D. subtitanus in having the short epig-
ynal teeth, anteriorly situated spermathecal heads and rounded, closely situated spermathecae but
can be distinguished by the epigynal teeth position (widely separated with atrium). The male is
similar to D. patellabifidus in having a bifurcate patellar apophysis and simple median apophysis
but can be distinguished by the slightly notched embolic base and the less apparent cymbial furrow
(the cymbial furrow might have been ignored in the original illustration).
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Female epigynal teeth small, widely separated; atrium small; copulatory ducts long, slender,
anteriorly extending; spermathecal heads situated anteriorly; spermathecae broad, rounded, close
together. Male palpal patellar apophysis strongly bifurcate, with ventral one large and dorsal one
small; RTA almost tibial length; lateral tibial apophysis present; cymbial furrow short; conductor
short, anteriorly extending, with small basal lamella; conductor dorsal apophysis short; embolus
broad, prolateral in origin; embolic base slightly notched; median apophysis long, broad, not
spoon-like.
DISTRIBUTION.— China (Tibet) (Map 11).
MATERIAL EXAMINED: CHINA: Tibet: Jala (SE Tibet), 29.42°N, 94.54°E, 2900 m., April 29,
1998, 3 females (G. Schaller, AMNH).
Draconarius hui (Dankittipakul and Wang, 2003), NEW COMBINATION
Map 12
Coelotes wangi Hu, 2001:133, figs. 844:1—2 (female holotype, 2 female paratypes from Sejilashan, Linzhi,
Tibet, China, in SDU, not examined). This specific name is preoccupied by Chen and Zhao, 1997.
Coelotes hui Dankittipakul and Wang, 2003:13. (replacement name).
WANG: EAST ASIA COELOTINE SPIDERS 535
DIAGNosIs.— The female of this species is similar to D. funiushanensis but can be distin-
guished by the widely separated spermathecal bases.
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Female with epigynal teeth situated near atrium; atrium large; copulatory ducts originating
posteriorly, extending mesad of spermathecae; spermathecal bases small, widely separated; sper-
mathecal bases broad, anteriorly expanded and converging. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 12).
MATERIAL EXAMINED.— None.
Draconarius huizhunesis (Wang and Xu, 1988), NEW COMBINATION
Map I1
Coelotes huizhunesis Wang and Xu, 1988:7, figs. 1-5 (female holotype, male and female paratypes from
Qiyun Mt., Aihui, China, in HTC, not examined).— Platnick, 2000-2002.
Coelotes huizhuneesis: Wang and Xu, 1988: 4 (invalid name, Platnick, 2000-2002).
Coelotes huizhouensis Song, Zhu and Chen, 1999:375, figs. 219O—P, 222E, 223H (invalid emendation,
Platnick, 2000-2002)
DIAGNOsIs.— The female of this species is similar to D. strophadatus in having long epigynal
teeth but can be distinguished by the widely separated and anteriorly converging spermathecal
stalks. The male is similar to D. ornatus in having a bifurcate conductor but can be recognized by
the slender embolus and the strongly elongated cymbial furrow.
DESCRIPTION.— See Wang and Xu (1988). Chelicerae with three promarginal, and two retro-
marginal teeth. Female with epigynal teeth situated anteriorly, close together, strongly elongated;
spermathecal bases small, close together; spermathecal stalks extending laterally, widely separat-
ed, convoluted, and anteriorly converging. Male palpal patellar apophysis present; RTA long; lat-
eral tibial apophysis widely separated from RTA; cymbial furrow short; conductor bifurcate; con-
ductor dorsal apophysis present; median apophysis spoon-like, rounded; embolic base narrow;
embolus long, posterior in origin.
DISTRIBUTION.— China (Aihu1) (Map 11).
MATERIAL EXAMINED.— None.
Draconarius incertus Wang, sp. nov.
Figures 35A—B; Map 12
Types.— Male holotype and 4 male paratypes from native forest in Gaoligongshan at 9.5 road
km ESE Pianma, 25°59’N, 98°40’E, el. 2500 m, Nujiang Prefecture, Yunnan, China (October
15-18, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (holotype male and 2
paratype males) and CAS (2 paratype males).
ETyMOLOGY.— The specific name refers to the uncertain generic placement because of the not
spoon-like median apophysis and the prolaterally originated embolus.
DIAGNOSIS.— The male of this new species is similar to D. dubius but can be recognized by
the short patellar apophysis (Figs. 35A—B).
MALE.— Total length 8.00. Carapace 4.00 long, 2.80 wide. Abdomen 4.00 long, 2.60 wide.
Eye sizes and interdistances: AME 0.14, ALE 0.20, PME 0.17, PLE 0.19; AME-AME 0.06, AME-
ALE 0.05, PME-PME 0.12, PME-PLE 0.15, AME-PME 0.12. Leg measurements: I: 17.7 (4.40,
5.70, 4.50, 3.10); I: 16.0 (4.00, 5.00, 4.10, 2.90); III: 14.2 (3.70, 4.30, 4.00, 2.20); IV: 19.1 (4.60,
536 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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5.56, 6.00, 2.90). Chelicerae with three promarginal and two retromarginal teeth. Male palp with
short patellar apophysis; RTA long; lateral tibial apophysis large, widely separated from RTA; cym-
bial furrow short; conductor slender, with apex slightly bifurcate; conductor dorsal apophysis slen-
der; conductor lamella small; embolus prolateral in origin; median apophysis large, not spoon-like
(Figs. 35A—B).
FEMALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 12).
OTHER MATERIAL EXAMINED.— None.
Draconarius infulatus (Wang et al., 1990)
Map 12
Coelotes infulatus Wang et al., 1990:202, figs. 57-58 (female holotype and female paratype from Tianmushan,
Zhejiang, China, in HBI, not examined).— Song, Zhu and Chen, 1999:375, figs. 220F—G.
Draconarius infulatus: Wang 2002:67.
DIAGNOsIS.— The female of this species can be easily distinguished from other coelotines by
the posteriorly situated, widely separated epigynal teeth and the broad, anteriorly expanded, over-
lapped copulatory ducts.
DESCRIPTION.— See Wang et al. (1990). Chelicerae with three promarginal, and two retromar-
ginal teeth. Female epigynal teeth widely separated, situated posteriorly near atrium; atrium broad,
near epigynal furrow; copulatory ducts broad, posteriorly originating, anteriorly expanded, con-
nected to spermathecae laterally; spermathecae small, enclosed by copulatory ducts. Male
unknown.
DISTRIBUTION.— China (Zhejiang) (Map 12).
MATERIAL EXAMINED.— None.
Draconarius jiangyongensis (Peng, Gong and Kim, 1996), NEW COMBINATION
Figures 36A—B; 96B; Map 12
Coelotes jiangyongensis Peng, Gong and Kim, 1996:19, figs. 7-9 (female holotype, 4 female paratypes from
Qianjiatong, Jiangyong, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:376, figs.
220J—K.
DIAGNOsIS.— This species can be easily recognized by the vase-shaped epigynal plate, the
long, anteriorly extending copulatory ducts, and the long, wedge-shaped spermathecae of female
(Fig. 36A—B).
DESCRIPTION.— See Peng, Gong and Kim (1996). Cheliceral promargin with three teeth,
retromargin four. Female epigynal teeth short, widely separated; atria separated by the median sep-
tum that spaced wider apart posteriorly than anteriorly; copulatory ducts long, anteriorly extending
along with spermathecae; spermathecal heads small, anteriorly situated and laterally extending;
spermathecal bases broad, widely separated; spermathecal stalks long, wider apart posteriorly and
converged together anteriorly (Figs. 36A—B). Male unknown.
DISTRIBUTION.— China (Hunan) (Map 12).
MATERIAL EXAMINED.— CHINA: Hunan: Jiangyong, Qianjiatong, October 1, 1991, female
holotype and 4 female paratypes (N.S. Gong, HBI).
Draconarius labiatus (Wang and Ono, 1998)
Figures. 37A—B; Map 13
WANG: EAST ASIA COELOTINE SPIDERS 537
Coelotes labiatus Wang and Ono, 1998:145, figs. 8-9 (female holotye and 1 female paratype from Nantou
Hsien, Taiwan, China, in NSMT, examined).
Draconarius labiatus: Wang, 2002:67.
DIAGNOsIS.— This species is similar to D. wenzhouensis but can be distinguished by the small,
dorsally covered spermathecal heads and the transversely extending spermathecal expansions (Fig.
37A-B).
DESCRIPTION.— See Wang and Ono (1998). Cheliceral promargin with three teeth, retromar-
gin with two. Female epigynal teeth short, anteriorly situated, adjacent; atrium situated posteriorly
near epigastric furrow, with anterior atrial margin expanded, lip-shaped; copulatory ducts slightly
sclerotized, originating posteriad, extending mesad of spermathecae; spermathecal heads small,
covered by copulatory ducts in dorsal view; spermathecal bases widely separated; spermathecal
stalks extending anteriorly, then curved and extending (Figs. 37A—B). Male unknown.
DISTRIBUTION.— China (Taiwan) (Map 13).
MATERIAL EXAMINED.— CHINA: Taiwan: Nantou Hsien, Tatachia, 2100 m alt., March 5, 1991,
female holotype (H. Ono, NSMT, NSMT-Ar.3436); Nantou Hsien, Shemu, Mt. Hohuan-shan, 2180
m alt., March 12, 1991, 1 female paratype (H. Ono, NSMT, NSMT-Ar.3444).
Draconarius linzhiensis (Hu, 2001), NEW COMBINATION
Map 13
Coelotes linzhiensis Hu, 2001:138, figs. 848:1—2 (female holotype and paratypes from Linzhi, Tibet, China,
in SDU, not examined).
DIAGNOsIS.— The female is similar to D. gingzangensis in having the similar spermathecal
tubes but can be distinguished by the slightly separated atria and the moderately expanded anteri-
or spermathecae (same width as stalks).
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Female epigynal teeth situated laterad of atrium; atrium large; spermathecal heads situated
medially on spermathecae; spermathecal bases widely separated; spermathecal stalks broad, ante-
riorly expanded and converging. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 13).
MATERIAL EXAMINED.— None.
Draconarius linxiaensis Wang, sp. nov.
Figures 38A—B; Map 13
TyprEs.— Male holotype from Linxia, Gansu, China (August 30, 1997; X.P. Wang), deposited
in IZB.
ETYMOLOGyY.— The specific name refers to the type locality.
DIAGNOsIS.— The male of this new species is similar to D. neixiangensis in lacking a patellar
apophysis and having a broad, deeply grooved conductor but can be distinguished by the broad
median apophysis and the slender embolic base and apex (Figs. 38A—B).
MALE.— Total length 7.80. Carapace 4.00 long, 2.72 wide. Abdomen 3.80 long, 2.00 wide.
Eye sizes and interdistances: AME 0.14, ALE 0.17, PME 0.15, PLE 0.17; AME-AME 0.06, AME-
ALE 0.05, PME-PME 0.13, PME-PLE 0.17, AME-PME 0.12. Leg measurements: I: 12.3 (3.20,
4.20, 2.92, 2.00); I: 11.3 (3.00, 3.80, 2.88, 1.60); III: 10.7 (2.80, 3.36, 3.00, 1.52); IV: 14.3 (3.20,
4.40, 4.32, 2.40). Cheliceral promargin with three teeth, retromargin with two. Male palp lacking
538 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
patellar apophysis; RTA long; lateral tibial apophysis large, widely separated from RTA; cymbial
furrow short; conductor broad, deeply grooved; conductor dorsal apophysis broad; embolus long,
broad, posterior in origin, with narrow base and slender, bifurcate apex; median apophysis broad,
spoon-like (Figs. 38A—B).
FEMALE.— Unknown.
DISTRIBUTION.— China (Gansu) (Map 13).
OTHER MATERIAL EXAMINED.— None.
Draconarius lutulentus (Wang et al., 1990)
Figures 39A—E; 40A—H; Map 13
Coelotes lutulentus Wang et al., 1990:216, figs. 88—92 (2 male and 2 female paratypes, holotype not indicat-
ed, from Zhangjiajie, Dayong, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:376, figs.
220R-S, 222K, 223N; -Hu, 2001:139, figs. 8-49:1-4.
Coelotes sinualis Chen, Zhao and Wang, 1991:10, figs. 3, 4 (female holotype from Jinding, Wudangshan,
Hubei, China, in HUW, examined);— Song, Zhu and Chen, 1999:378, figs. 224T, U. NEW SYNONYMY.
Draconarius sinualis: Wang, 2002:72, figs. 192—210.
Draconarius lutulentus: Wang, 2002:12.
DIAGNOsIS.— The female of this species is similar to D. wuermilii but can be distinguished by
the strongly convoluted spermathecae and the differences in shapes between their atria (Figs.
39A-B; 40A—B, E-F). The male is similar to D. wudangensis in having the short RTA but can be
recognized by the long, slender conductor (Figs. 39C—E; 40C—D, G-—H).
SYNONYMY.— This species is widespread in central and eastern China, with variable male and
female genitalic morphology. The species D. sinualis, described from Wudangshan, Hubei, China,
has the identical genitalic morphology and is placed as a junior synonym of D. lutulentus.
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin three. Epigynal teeth short, situated laterally, near atrium; atrium small; copulatory ducts
long, originating posteriorly, extending mesad of spermathecae, looped around distal spermathe-
cae; spermathecal heads small; spermathecal bases widely separated; spermathecal stalks long,
anteriorly converging (Figs. 39A—B; 40A—B, E—F). Male palp with large patellar apophysis; RTA
short, approximately half tibial length; lateral tibial apophysis large; cymbial furrow long, about
half tibial length or longer; conductor long, slender, with large basal lamella; conductor dorsal
apophysis slender; embolus posterior in origin; median apophysis spoon-like, elongated (Figs.
39C-E; 40C—D, G-H).
DISTRIBUTION.— China (Hubei, Hunan, Shaanxi, Tibet, Zhejiang, Anhui) (Map 13).
MATERIAL EXAMINED.— CHINA: Hubei: Wudangshan, Jinding, April 23, 1982, female holo-
type of Coelotes sinualis Chen, Zhao and Wang, 1991 (HUW); Wudangshan, Jinding, September
24, 1997, 3 females, 1 male and 2 females (X.P. Wang, AMNH); Wudangshan, Nanya to Jinding,
September 24, 1997, 1 male and 1 female, 1 female, 11 females and 3 males (X.P. Wang, IZB);
Wudangshan, Zhixiao to Nanya, September 23, 1997, 3 females and 7 males (X.P. Wang, MCB);
Hongping, September 21, 1997, 2 males and 9 females (X.P. Wang, IZB). Hunan: Dayong,
Zhangjiajie, October 17, 1984, 2 male and 2 female paratypes (J.F. Wang and Y.J. Zhang, HBI).
Shaanxi: Huxian, Cuihuashan, October 18, 1989, 1 male and | female (X.P. Wang, IZB).
Draconarius magniceps (Schenkel, 1936), NEW COMBINATION
Figure 41A—B; Map 14
WANG: EAST ASIA COELOTINE SPIDERS 539
Coelotes magniceps Schenkel, 1936:186, fig. 61 (female holotype from Kina, S. Gansu, China, in NRS, exam-
ined).— Song, Zhu and Chen, 1999:376.
DrAGNosis.— The female of this species can be easily recognized by the absence of epigynal
teeth, the presence of a broad atrial septum, and the large, anteriorly expanded copulatory ducts
(Figs. 41A—B).
DESCRIPTION.— Described by Schenkel (1936). Chelicerae with three promarginal and two
retromarginal teeth. Female epigynal teeth absent; atrium large, with broad septum; copulatory
ducts broad, extending anteriorly; spermathecal heads small, anteriorly situated; spermathecae
broad, short, slightly separated (Figs. 41A—B). Male unknown.
DISTRIBUTION.— China (Gansu) (Map 14).
MATERIAL EXAMINED.— CHINA: Gansu: Kina, female holotype (Dr. Hummel, NRS, burk 6,
Sven Hedins Exp. Ctr. Asien).
Draconarius molluscus (Wang et al., 1990)
Figure 42A-B; 96D; Map 14
Coelotes molluscus Wang et al., 1990:214, figs. 86-87 (4 female types, holotype not indicated, from Lushan,
Jiangxi, China, in HBI, examined).— Song, Zhu and Chen, 1999:376, figs. 221G—H.
Draconarius molluscus: Wang, 2002:67.
DIAGNOSIS.— The female of this species is similar to D. haopingensis in lacking the epigynal
teeth and having the laterally situated epigynal hoods but can be distinguished by the presence of
two copulatory duct loops and the convoluted spermathecal distal ends (Fig. 42A—B).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin three. Female without epigynal teeth; atrium broad; epigynal hoods deep, situated posteri-
orly, laterad of atrium; copulatory ducts long, originating posteriorly, mesad of spermathecae,
looped around spermathecae; spermathecal heads small; spermathecal bases widely separated;
spermathecal stalks long, anteriorly converging (Figs. 42A—B). Male unknown.
DISTRIBUTION.— China (Jiangxi) (Map 14).
MATERIAL EXAMINED. —CHINA: Jiangxi: Lushan, June 15, 1987, 4 female types (J.F. Wang,
HBI).
Draconarius nanyuensis (Peng and Yin, 1998), NEw COMBINATION
Figures 43A—B; Map 14
Coelotes nanyuensis Peng and Yin, 1998:27, figs. 7-9 (female holotype from Nanyue, Hunan, China, in HBI,
examined).— Song, Zhu and Chen, 1999:376.
DIAGNOSIS.— The female of this species can be easily distinguished by the adjacent, anterior-
ly situated epigynal teeth, the separated atria, and the rounded spermathecae (Figs. 43A—B).
DESCRIPTION.— Described by Peng and Yin (1998). Promargin of chelicera with three teeth,
retromargin with two. Female epigynal teeth short, situated anteriorly, adjacent; atria small, dis-
tinctly separated; copulatory ducts narrow, originating posteriorly, extending mesad of spermathe-
cae; spermathecal heads anteriorly situated; spermathecae large, rounded, slightly separated (Figs.
43A—B). Male unknown.
DISTRIBUTION.— China (Hunan) (Map 14).
MATERIAL EXAMINED.— CHINA: Hunan: Nanyue, August 3—7, 1995, female holotype (C.M.
Yin, HBI).
540 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconarius neixiangensis (Hu, Wang and Wang, 1991)
Figures 44A—D; Map 14
Coelotes neixiangensis Hu, Wang and Wang, 1991:43, figs. 18-21 (female holotype, male and female
paratypes from Neixiang, Henan, China, deposited in SDU, not examined).— Song, Zhu and Chen,
1999:377, figs. 221K—L, 222N, 224B.
Coelotes baccatus Wang, 1994:286, figs. 1-5 (2 male and 2 female types from Xiangfan, Hubei, China, in
HBI, examined).— Song, Zhu and Chen, 1999:374, figs. 216P—Q, 218E, L. NEW SYNONYMY.
Draconarius baccatus: Wang, 2002:66.
Draconarius neixiangensis: Wang, 2002:68.
DIAGNOsIS.— The female of this species can be easily recognized by the anteriorly situated
epigynal teeth, the broad, posteriorly situated atrium, and the broad, anteriorly expanded copulato-
ry ducts (Figs. 44A—B). The male is similar to D. linxiaensis in lacking a patellar apophysis and
having the broad, deeply grooved conductor but can be distinguished by the elongated, slender
median apophysis and the broad embolic base and apex (Figs. 44C—D).
DESCRIPTION.— Described by Chen (1984) and Wang (1994). Chelicerae with three promar-
ginal and two retromarginal teeth. Female epigynal teeth short, situated anteriorly, close together;
atrium broad, near epigastric furrow, with anterior margin slightly notched into two broad, less
sclerotized pieces; copulatory ducts broad, originating posteriorly, strongly expanded anteriorly;
spermathecal heads small, situated laterally on ventral side of spermathecae; spermathecal bases
small, widely separated; spermathecal stalks short, anteriorly diverging (Figs. 44A—B). Male palp
without patellar apophysis; RTA long; lateral tibial apophysis widely separated from RTA; cymbial
furrow short; conductor broad, strongly grooved, with small basal lamella; conductor dorsal apoph-
ysis small; embolus broad, posterior in origin, with slightly modified apex; median apophysis
spoon-like, elongated (Figs. 44C—D).
DISTRIBUTION.— China (Henan, Hubei) (Map 14).
MATERIAL EXAMINED.— CHINA: Hubei: Xiangfan, October 26, 1990, 2 male and 2 female
types of Coelotes baccatus (J.F. Wang, HBI).
Draconarius nudulus Wang, sp. nov.
Figures 45A—B; Map 15
Types.— Male holotype from 36 air km SE TengChong, pass over Gaoligongshan at 2100 m,
24°50'N, 98°47’E, Nankang, Baoshan, Yunnan, China (November 4-7, 1998; C. Griswold, D.
Kavanaugh, C-L. Long), deposited in HBI.
ETYMOLOGy.— The specific name refers to the absence of patellar apophysis.
DIAGNOsISs.— The male of this new species is similar to D. argenteus in lacking a patellar
apophysis and having a short conductor but can be distinguished by the long RTA (more than half
tibial length), the simple median apophysis, and the non-lobed embolic base (Figs. 45A—B).
MALE.— Total length 6.14. Carapace 3.14 long, 1.94 wide. Abdomen 3.00 long, 2.30 wide.
Eye sizes and interdistances: AME 0.08, ALE 0.16, PME 0.15, PLE 0.15; AME-AME 0.04, AME-
ALE 0.04, PME-PME 0.15, PME-PLE 0.15, AME-PME 0.10. Leg measurements: I: 8.68 (2.40,
3.08, 1.96, 1.24); I: 7.36 (2.08, 2.40, 1.76, 1.12); III: 6.70 (1.86, 2.06, 1.80, 0.98); IV: 9.30 (2.46,
3.00, 2.64, 1.20). Promargin of chelicera with three teeth, retromargin one. Male palp without
patellar apophysis; RTA approximately tibial length or slightly longer; lateral tibial apophysis near
RTA; cymbial furrow short; conductor short; conductor dorsal apophysis broad; conductor lamel-
la small; embolus posterior in origin; median apophysis strongly elongated, slightly or not spoon-
like (Figs. 45A—B). Female unknown.
WANG: EAST ASIA COELOTINE SPIDERS 54]
FEMALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 15).
OTHER MATERIAL EXAMINED.— None.
Draconarius ornatus (Wang et al., 1990), NEW COMBINATION
Figures 46A—C; 96C; Map 15
Coelotes ornatus Wang et al., 1990:199, figs. 53-54 (female holotype and paratypes from Xishan, Kunming,
Yunnan, China, in HBI, not examined).— Song, Zhu and Chen, 1999:377, figs. 2210-P.
DIAGNosIs.— The female of this species is similar to D. terebratus by lacking epigynal teeth,
having posteriorly situated epigynal hoods, anteriorly extending copulatory ducts, and long sper-
mathecal heads but can be distinguished by the slightly elongated atrium and the less broad sper-
mathecal bases. The male is similar to D. huizhunesis by having a bifurcate conductor but can be
distinguished by the broad embolus and the short cymbial furrow (Fig. 46A—C).
Nores.— Although the female types could not be located, two males labeled as C. ornatus
were found in HBI collection. Further collection of this species is badly needed in order to verify
this association.
DESCRIPTION.— The female was described by Wang et al. (1990), and the male is described
for the first time. Cheliceral promargin with three teeth, retromargin with two. Female epigynal
teeth absent; atrium large; epigynal hoods situated posteriorly, laterad of atrium; copulatory ducts
posteriorly originating, strongly expanded anteriorly; spermathecal heads long, slender, anteriorly
extending: spermathecal bases small, widely separated; spermathecal stalks anteriorly diverging.
Male palp with patellar apophysis long; RTA small; lateral tibial apophysis widely separated from
RTA; cymbial furrow short; conductor long, broad, with bifurcate apex and small basal lamella;
conductor dorsal apophysis small; embolus broad, posterior in origin; median apophysis small,
spoon-like (Figs. 46A-C).
DISTRIBUTION.— China (Yunnan) (Map 15).
MATERIAL EXAMINED.— CHINA: Yunnan: Kunming, Xishan, August 8, 1991, 2 males (HBI).
Draconarius parabrunneus Wang, sp. nov.
Figures 47A—B; 96E; Map 15
TyprEs.— Female holotype, 4 female paratypes from native forest in Gaoligongshan at 9.5 road
km ESE Pianma, 25°59’N, 98°40’E, el. 2500 m, Gaoligongshan, Nujiang Prefecture, Yunnan,
China (October 15—18, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (holotype
and 2 paratype females) and CAS (2 paratype females).
ETYMOLOGyY.— The specific name refers to its similarity to D. brunneus related species.
DIAGNOsIS.— The female of this species is similar to D. pseudobrunneus in lacking the epig-
ynal teeth and having small, broad spermathecae but can be distinguished by the anteriorly situat-
ed spermathecal heads and the broad spermathecal bases (broader than stalks) (Figs. 47A—B).
FEMALE.— Total length 10.3. Carapace 4.30 long, 2.80 wide. Abdomen 6.00 long, 4.00 wide.
Eye sizes and interdistances: AME 0.13, ALE 0.22, PME 0.20, PLE 0.20; AME-AME 0.10, AME-
ALE 0.04, PME-PME 0.13, PME-PLE 0.13, AME-PME 0.14. Leg measurements: I: 12.8 (3.60,
4.30, 2.90, 2.00); II: 11.6 (3.30, 3.80, 2.76, 1.70); III: 10.6 (3.00, 3.20, 2.80, 1.60); IV: 14.3 (3.60,
4.56, 4.10, 2.00). Promargin of chelicera with three teeth, retromargin with two. Female epigynal
teeth absent; atrium small; epigynum wrinkly on anterior atrium; copulatory ducts short, situated
542 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
mesad of spermathecae; spermathecal heads situated anteriorly; spermathecal bases broad, widely
separated; speramthecal stalks short (Figs. 47A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 15).
OTHER MATERIAL EXAMINED.— None.
Draconarius paraterebratus Wang, sp. nov.
Figures 48A—B; Map 16
Types.— Female holotype from pass over Gaoligongshan at 2300 m, Luoshuidong, 28 air km
E TengChong, 24°57’N, 98°45’E, native forest, TengChong, Luoshuidong, Baoshan, Yunnan,
China (October 26-31, 1998; C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI.
ETYMOLOGY.— The specific name refers to its similarity to D. terebratus.
DIAGNOsIS.— The female of this species can be easily recognized by the posteriorly situated
epigynal teeth and the presence of diverticula on spermathecal bases (Figs. 48A—B).
FEMALE.— Total length 11.2. Carapace 5.00 long, 3.20 wide. Abdomen 6.20 long, 4.20 wide.
Eye sizes and interdistances: AME 0.25, ALE 0.24, PME 0.24, PLE 0.24; AME-AME 0.10, AME-
ALE 0.10, PME-PME 0.20, PME-PLE 0.26, AME-PME 0.25. Leg measurements: I: 13.5 (3.82,
4.64; 3.28, 1.80); W: 11-9 G.50; 4.02, 2.82, 1.58); MI: 10:3 (3:04, 3.22, 2:76, 1-30): ANE 1S Grs2
4.32, 3.70, 1.70). Promargin of chelicera with three teeth, retromargin with two. Female epigynal
teeth small, situated posteriorly laterad of atrium; atrium broad; copulatory ducts broad, long, orig-
inating anteriorly, overlapped dorsally with spermathecal stalks; spermathecal heads anteriorly sit-
uated; spermathecal bases slightly separated, with long diverticula; spermathecal stalks long, wide-
ly separated (Figs. 48A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Fig. Map 16).
OTHER MATERIAL EXAMINED.— None.
Draconarius patellabifidus Wang, sp. nov.
Figures 49A—D; Map 17
Types.— Male holotype, 2 male and 4 female paratypes from Dulong/Gongshan Yakou area,
Nujiang State Nature Reserve, Nujiang Prefecture, Gaoligong Shan, Yunnan, China (July 16-17,
2000; H.M. Yan, D. Kavanaugh, C.E. Griswold, H.B. Liang, D. Ubick, and D.Z. Dong), deposited
in HBI (male holotype, 1 male and 3 female paratype) and CAS (1 male and | female paratypes);
1 female paratype from Nujiang State Nature Reserve, Dulong/Gongshan Yakou area, 21 airkm W
of Gongshan, N27.697°/E98.454°, 3300-3680m, Gaoligong Shan, Nujiang Prefecture, Yunnan,
China (July 16-17, 2000; H.M. Yan, D. Kavanaugh, C.E. Griswold, H.B. Liang, D. Ubick, and
D.Z. Dong), deposited in CAS; 9 male paratypes from Pianma, Yakou, Pass over Gaoligongshan at
el. 3200 m, 25°58’N, 98°41’E, bamboo thicket and under stones, Nujiang Prefecture, Yunnan,
China (October 11, 1998: C. Griswold, D. Kavanaugh, C-L. Long), deposited in HBI (4 males) and
CAS (5 males).
ETYMOLOGyY.— The specific name refers to the bifurcate patellar apophysis.
DIAGNOsIs.— The female of this species is similar to D. dubius and D. simplicidens by lack-
ing epigynal teeth and having broad spermathecae but can be recognized from D. simplicidens by
the anteriorly situated spermathecal heads, from D. dubius by the less expanded spermathecal bases
(smaller than stalks) (Figs. 49A—B). The male is similar to D. himalayaensis by having the bifur-
WANG: EAST ASIA COELOTINE SPIDERS 543
cate patellar apophysis and the simple median apophysis but can be distinguished by the entire
embolic base and the presence of distinct cymbial furrow (Figs. 49C—D).
FEMALE.— Total length 10.4. Carapace 5.20 long, 3.51 wide. Abdomen 5.20 long, 3.38 wide.
Eye sizes and interdistances: AME 0.15, ALE 0.25, PME 0.20, PLE 0.20; AME-AME 0.13, AME-
ALE 0.10, PME-PME 0.15, PME-PLE 0.25, AME-PME 0.23. Leg measurements: I: 13.2 (3.64,
Beyer -0)/ 51-95); Me 12-21(3:385 4.16; 2-81, 1-82); MMe 115 GRO2s 31855122945. 1-69): IV14 7 C.77,
4.89, 3.98, 2.05). Promargin of chelicera with three teeth, retromargin with two. Epigynal teeth
absent; atrium small; copulatory ducts small, originating posteriorly, extending mesad of spermath-
ecae; spermathecal heads situated anteriorly; spermathecae broad, widely separated (Figs. 49A—B).
MALeE.— Total length 11.8. Carapace 6.11 long, 4.21 wide. Abdomen 5.72 long, 3.64 wide.
Eye sizes and interdistances: AME 0.19, ALE 0.26, PME 0.23, PLE 0.24; AME-AME 0.11, AME-
ALE 0.06, PME-PME 0.15, PME-PLE 0.20, AME-PME 0.18. Leg measurements: I: 21.5 (5.51,
One 5-59, 3:28): We 19.8 6.20; 6:50; 5.20; 2:91): Ml: 17-6 (4:5595:59;.4.81; 2.60); IV: 21-7. (59,
6.76, 6.24, 3.07). Promargin of chelicera with three teeth, retromargin with two. Palpal patellar
apophysis strongly bifurcate, with ventral one large and dorsal one small; RTA long; lateral tibial
apophysis wide apart from RTA; cymbial furrow short; conductor short, anteriorly extending, with
bifid apex and small basal lamella; conductor dorsal apophysis short; embolus broad, prolateral in
origin; median apophysis long, not spoon-like (Figs. 49C—D).
DISTRIBUTION.— China (Yunnan) (Map 17).
OTHER MATERIAL EXAMINED.— None.
Draconarius penicillatus (Wang et al., 1990), NEW COMBINATION
Figures 50A—E; Map 16
Coelotes penicillatus Wang et al., 1990:197, figs. 48-52 (female holotype and male paratype from Xishan,
Kunming, Yunnan, China, in HBI, examined).
Coelotes penicilatus Song, Zhu and Chen, 1999:377, figs. 221U-V, 223A, 224E.
DiIAGNOsIs.— The female of this species can be easily identified by the anteriorly situated cop-
ulatory ducts and the laterally extending spermathecae (Fig. 50A—B). The male is similar to D.
coreanus but can be recognized by the posteriorly extending conductor and the short embolic base
(Figs. 50C-E).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth small, situated near atrium; atrium small; copulatory ducts
broad, anteriorly situated, convoluted around spermathecal heads; spermathecal heads anteriorly
Situated; spermathecal bases small; spermathecal stalks extending laterally (Figs. S0A—B). Male
plalp with small patellar apophysis; RTA long; lateral tibial apophysis large; cymbial furrow more
than half cymbial length; conductor short, posteriorly extending, with small basal lamella; conduc-
tor dorsal apophysis broad; embolus long, posterior in origin; embolic base short; median apoph-
ysis small, rounded; spoon-like (Figs. 50C-E).
DISTRIBUTION.— China (Yunnan) (Map 16).
MATERIAL EXAMINED.— CHINA: Yunnan: Kunming, Xishan, October 24, 1987, female holo-
type and male paratype (J. F Wang, HBI).
Draconarius pervicax (Hu and Li, 1987), NEw COMBINATION
Map 15
Coelotes pervicax Hu and Li, 1987:279, figs. 18.5-6 (female holotype from Yadong, Tibet, China, in SDU,
not examined).— Song, Zhu and Chen, 1999:377, figs. 221W-X;— Hu, 2001:141, figs. 8-51:1-2.
544 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
DIAGNOSIS.— The female of this species is similar to D. quadratus but can be distinguished
by the closely situated spermathecal bases.
DESCRIPTION.— See Hu and Li (1987). Chelicerae with three promarginal, and two retromar-
ginal teeth. Female epigynal teeth situated laterad of atrium; atrium broad; spermathecal bases
large, close together; spermathecal stalks widely separated, anteriorly elongated. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 15).
MATERIAL EXAMINED.— None.
Draconarius picta (Hu, 2001), NEW COMBINATION
Map 16
Coelotes picta Hu, 2001:142, figs. 8-52:1—2 (female holotype and 2 female paratypes from Sejilashan, Linzhi,
Tibet, China, in SDU, not examined).
DIAGNOSIS.— The female is similar to D. wudangensis in having the medially situated and
widely separated epigynal teeth, and similar spermathecal tubes but can be distinguished by the
closely situated spermathecae and the slender spermathecal bases.
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Female epigynal teeth widely separated, situated posteriorly near epigastric furrow; atrium
small; spermathecal heads small; spermathecal bases slightly separated; spermathecal stalks broad,
anteriorly expanded and converging. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 16).
MATERIAL EXAMINED.— None.
Draconarius potanini (Schenkel, 1963)
Figures 51A—B; Map 16
Cybaeus potanini Schenkel, 1963:275, fig. 156 (female holotype from Gansu, China, in MNHN, exam-
ined).— Song, Zhu and Chen, 1999:355, figs. 208I-J.
Draconarius potanini: Wang, 2002:68.
DIAGNOsIs.— The female of this species is similar to D. baxiantaiensis by having the looped
copulatory ducts and broad, long spermathecae but can be distinguished by the small atrium, the
presence of two copulatory duct loops, and the anteriorly situated spermathecal heads (Figs.
S51A-B).
DESCRIPTION.— Described by Schenkel (1963). Cheliceral promargin with three teeth, retro-
margin with two. Female with epigynal teeth small; atrium small; copulatory ducts long, originat-
ing posteriorly, with two loops around spermathecae; spermathecal heads small, situated ventrally
on anterior spermathecae; spermathecal bases broad, widely separated; spermathecal stalks broad,
anteriorly converging (Figs. 51 A—B). Male unknown.
DISTRIBUTION.— China (Gansu) (Map 16).
MATERIAL EXAMINED.— CHINA: Gansu: Kloster Dschoni (Choni), June 8, 1885, female holo-
type (MNHN).
Draconarius pseudobrunneus Wang, sp. nov.
Figure 52A—B; Map 18
Typrs.— Female holotype and 3 female paratypes from Danzhu He drainage, 13.5 air km
WANG: EAST ASIA COELOTINE SPIDERS 545
SSW of Gongshan, 2700m, N27.631°/E98.621°, Gongshan Co., Nujiang, Yunnan, China (June 30—
July 5, 2000; D. Kavanaugh, C.E. Griswold, H.B. Liang, D. Ubick, H M. Yan, and D.Z. Dong),
deposited in HBI (holotype female and | paratype female) and CAS (2 paratype females).
EryMoLoGy.— The specific name refers to its similarity to D. brunneus.
DIAGNosis.— The female of this species is similar to D. parabrunneus in lacking the epigy-
nal teeth, and having small, broad spermathecae but can be distinguished by the medially situated
spermathecal heads and the moderately expanded spermathecal bases (same width as stalks) (Figs.
52A-B).
FEMALE.— Total length 7.28. Carapace 3.38 long, 2.08 wide. Abdomen 3.90 long, 2.73 wide.
Eye sizes and interdistances: AME 0.09, ALE 0.18, PME 0.15, PLE 0.17; AME-AME 0.09, AME-
ALE 0.05, PME-PME 0.10, PME-PLE 0.15, AME-PME 0.13. Leg measurements: I: 7.72 (2.31,
MoUG4, 1217); I:-7.26 2.08, 2:55, 1-59, 1.04); Ml: 6.79 (1-87, 2.24; 1.69;,0.99): TV: 9.8112:42,
3.07, 2.47, 1.22). Promargin of chelicera with three teeth, retromargin with two. Epigynal teeth
absent; atrium small; epigynum wrinkled on anterior atrium; copulatory ducts small, situated
mesad of spermathecae; spermathecal heads small, situated medially on spermathecae; spermathe-
cal bases broad, widely separated; spermathecal stalks widely separated, slightly converging ante-
riorly (Figs. 52A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 18).
OTHER MATERIAL EXAMINED.— None.
Draconarius pseudocapitulatus Wang, sp. nov.
Figures 53A—B; Map 19
Typrs.— Female holotype and | female paratype from Danzhu He drainage, 13.5 air km SSW
of Gongshan, 2700m, N27.631°/E98.621°, Gongshan Co., Nujiang, Yunnan, China (June 30 — July
5, 2000; D. Kavanaugh, C.E. Griswold, H.B. Liang, D. Ubick, H.M. Yan, and D.Z. Dong), deposit-
ed in HBI; | female paratype from Nujiang State Nature Reserve, No. 12 Bridge Camp area, 16.3
air km W of Gongshan, N27.715°/E98.502°, 2775m, Nujiang Prefecture, Gaoligong Shan, Yunnan,
China (July 15—19, 2000; H.M. Yan, D. Kavanaugh, C.E. Griswold, H.B. Liang, D. Ubick, and D.
Z. Dong), deposited in CAS.
ETYMOLOGyY.— The specific name refers to its similarity to D. capitulatus.
DiAGNosis.— The female of this species is similar to D. capitulatus but can be distinguished
by the anteriorly expanded, widely separated spermathecae (fig. 53A—B).
FEMALE.— Total length 11.7. Carapace 5.20 long, 3.54 wide. Abdomen 6.50 long, 4.94 wide.
Eye sizes and interdistances: AME 0.15, ALE 0.23, PME 0.20, PLE 0.22; AME-AME 0.14, AME-
ALE 0.11, PME-PME 0.19, PME-PLE 0.29, AME-PME 0.20. Leg measurements: I: 12.0 (3.64,
ow 2/0, 1.69): Ie 11-7 G25, 4.50; 2.47, 1.46); WI: 9:60 @.73; 3.15, 2:42, 1.30); IV: 13:26-64,
4.42, 3.51, 1.61). Promargin of chelicera with three teeth, retromargin with two. Epigynal teeth
absent; atrium broad, near epigastric furrow; copulatory ducts short, situated mesad of spermathe-
cae; spermathecal heads situated anteriorly, mesad of spermathecae; spermathecal bases broad,
widely separated; spermathecal stalks short, expanded anteriorly (Figs. 53A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 19).
OTHER MATERIAL EXAMINED.— None.
546 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconarius pseudowuermlii Wang, sp. nov.
Figures 54A—B; Map 18
TyprEs.— Female holotype from Pass over Gaoligongshan at 2300 m, Luoshuidong, 28 air km
E TengChong, 24°57’N, 98°45’E, native forest, Baoshan, Yunnan, China (October 26-31, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI.
ETYMOLOGY.— The specific name refers to its similarity to D. wuermilii.
DiAGNosis.— The female of this new species is similar to D. wuermlii but can be distin-
guished by the medially situated, not looped copulatory ducts and the broad, rounded spermathe-
cal bases (fig. 54A—B).
FEMALE.— Total length 6.30. Carapace 3.10 long, 2.20 wide. Abdomen 3.20 long, 2.40 wide.
Eye sizes and interdistances: AME 0.10, ALE 0.18, PME 0.18, PLE 0.20; AME-AME 0.05, AME-
ALE 0.08, PME-PME 0.04, PME-PLE 0.13, AME-PME 0.12. Leg measurements: I: 7.44 (2.24,
2.56, 1.66, 0.98); I: 6.38 (1.96, 2.16, 1.46, 0.80); III: 5.32 (1.56, 1.80, 1.28, 0.68); IV: 7.64 (2.24,
2.64, 1.92, 0.84). Promargin of chelicera with three teeth, retromargin with two. Epigynal teeth
short, situated laterad of atrium; atrium broad, with less sclerotized, whitish median piece; copula-
tory ducts broad, anteriorly extending, situated mesad of spermathecae, connected to spermathecae
laterally; spermathecal heads large, widely separated; spermathecal bases broad, rounded; sper-
mathecal stalks long, strongly convoluted (Figs. 54A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 18).
OTHER MATERIAL EXAMINED.— None.
Draconarius qingzangensis (Hu, 2001), NEW COMBINATION
Map 17
Coelotes gingzangensis Hu, 2001:143, figs. 8-54:1—2 (female holotype, 2 female paratypes from Nanggian,
Qinghai, China, in SDU, not examined).
DIAGNosis.— The female is similar to D. linzhiensis by the similar spermathecal tubes but can
be distinguished by the not separated atria and the strongly expanded anterior spermathecae (larg-
er then stalks).
DESCRIPTION.— See Hu (2001). Chelicerae with three promarginal, and two retromarginal
teeth. Female epigynal teeth situated laterad of atrium, near atrial lateral margins; atrium large;
spermathecal heads situated medially on spermathecae; spermathecal bases widely separated; sper-
mathecal stalks broad, anteriorly expanded and converging. Male unknown.
DISTRIBUTION.— China (Qinghai) (Map 17).
MATERIAL EXAMINED.— None.
Draconarius quadratus (Wang et al., 1990), NEW COMBINATION
Figures 55A—B; Map 17
Coelotes quadratus Wang et al., 1990:197, figs. 46-47 (female holotype from Damingshan, Guangxi, China,
in HBI, examined).— Song, Zhu and Chen, 1999:377, figs. 224H-I.
DIAGNOsIS.— The female of this species is similar to D. pervicax but can be distinguished by
the widely separated spermathecal bases and the posteriorly situated epigynal hoods (Fig. 55sA—B).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth small, situated near atrium; atrium small; epigynal hoods
WANG: EAST ASIA COELOTINE SPIDERS 547
situated posteriorly near epigastric furrow; copulatory ducts small; spermathecal heads anteriorly
situated; spermathecal bases widely separated, broad; spermathecal stalks broad, anteriorly diverg-
ing (Figs. 55A—B). Male unknown.
DISTRIBUTION.— China (Guangxi) (Map 17).
MATERIAL EXAMINED.— CHINA: Guangxi: Damingshan, August 10, 1982, female holotype
(J.-F. Wang, HBI).
Draconarius rotundus Wang, sp. nov.
Figures 56A—B; Map 17
Types.— Female holotype from Pass over Gaoligongshan at 2300 m, Luoshuidong, 28 air km
E TengChong, 24°57’N, 98°45’E, native forest, Baoshan, Yunnan, China (October 26-31, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI.
ETYMOLOGyY.— The specific name refers to the rounded copulatory ducts.
DIAGNOsIS.— The female of this new species can be easily recognized by the posteriorly sit-
uated epigynal hoods, the anteriorly expanded copulatory ducts, and the looped, widely separated
spermathecae (Figs. 56A—B).
FEMALE.— Total length 8.00. Carapace 3.80 long, 2.60 wide. Abdomen 4.20 long, 2.80 wide.
Eye sizes and interdistances: AME 0.12, ALE 0.18, PME 0.19, PLE 0.19; AME-AME 0.09, AME-
ALE 0.10, PME-PME 0.05, PME-PLE 0.20, AME-PME 0.16. Leg measurements: I: 8.78 (2.62,
3.00, 1.92, 1.24); Il: 8.24 (2.42, 2.90, 1.76, 1.16); II: 6.74 (1.96, 2.18, 1.60, 1.00); IV: 10.6 (2.80,
3.38, 3.30, 1.16). Promargin of chelicera with three teeth, retromargin with two. Epigynal teeth
short, situated laterally, slightly anterad of atrium; atrium broad; epigynal hoods situated posterior-
ly laterad of atrium; copulatory ducts broad, originating posteriorly, mesad of spermathecae,
extending anteriorly and connected to spermathecae laterally; spermathecal heads situated lateral-
ly; spermathecal bases broad, widely separated; spermathecal stalks broad, looped (Figs. 56A—B).
MALE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 17).
OTHER MATERIAL EXAMINED.— None.
Draconarius rufulus (Wang et al., 1990), NEW COMBINATION
Figures 57A—E; 961; Map 18
Coelotes rufulus Wang et al., 1990:194, figs. 41-45 (2 male and 2 female types, holotype not indicated, from
Tianmushan, Zhejiang, China, in HBI, examined).— Song, Zhu and Chen, 1999:377, figs. 224L—M, 226S,
228A.
Coelotes rufuloides Zhang, Peng and Kim, 1997:295, figs. 8-9 (female holotype from Tiantong Mt., Zhejiang,
China, in HBI, examined). NEw SYNONYMY.
DIAGNOsIS.— This species can be easily recognized by the absence of epigynal teeth, the
tongue-like posterior epigynal extension, the elongated, looped copulatory ducts of female (Figs.
57A-B) and by lacking a patellar apophysis, the strongly modified conductor, and the broad,
strongly modified embolus of male (Figs. 57C-E). |
DESCRIPTION.— Described by Wang et al. (1990). Chelicerae with three promarginal and two
retromarginal teeth. Female epigynum without epigynal teeth; atrium small, situated posteriorly,
near epigastric furrow; copulatory ducts posteriorly originating, extending mesad of spermathecae,
and then strongly looped laterad of spermathecae; spermathecal heads small; spermathecal bases
widely separated; spermathecal stalks strongly convoluted, anteriorly elongated (Figs. 57A—B).
548 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Male palp without patellar apophysis; RTA long; lateral tibial apophysis large, near RTA; cymbial
furrow slightly shorter than half cymbial length; conductor long, broad, posteriorly extending,
strongly modified with broad dorsal edge and a strong tooth, with small basal lamella; conductor
dorsal apophysis small; embolus long, broad, strongly modified, posterior in origin; median apoph-
ysis spoon-like, elongated (Figs. 57C-—E).
DISTRIBUTION.— China (Anhui, Zhejiang) (Map 18).
MATERIAL EXAMINED.— CHINA: Zhejiang: Tianmushan, October 15, 1974, 2 male and 2
female paratypes (J.F. Wang, HB); Tiantong Mt., January 22, 1988, female holotype of Coelotes
rufuloides (Y.J. Zhang, HBI); Beihai Hotel (no provinces on label), October 29, 1974, 2 females
(C. D. Zhu, NBUMS, 74-1990).
Draconarius schenkeli (Brignoli, 1978)
Figures 58A—B; Map 18
Coelotes schenkeli Brignoli, 1978: 46, figs. 23-24 (female holotype from Chimakothi, Bhutan, in NHMB,
examined).
Draconarius schenkeli: Wang, 2002:69.
DIAGNOsIS.— The female of this species can be easily identified by the closely situated epig-
ynal teeth, the posteriorly situated epigynal hoods, the long, anteriorly extending copulatory ducts,
and the laterally situated spermathecal heads (Figs. 58A—B).
DESCRIPTION.— Described by Brignoli (1978). Promargin of chelicera with three teeth, retro-
margin with two. Female epigynal teeth short, situated medially on anterior atrial margin; epigynal
hoods situated posteriorly, laterad of atrium; copulatory ducts strongly extending and converging
anteriorly; spermathecal heads small, laterally situated; spermathecal bases small, slightly separat-
ed; spermathecal stalks short, slightly extending laterally (Figs. 583A—B). Male unknown.
DISTRIBUTION.— Bhutan (Map 18).
MATERIAL EXAMINED.— BHUTAN: Chimakothi, 1900-2300 m, May 22, 1972, female holotype
(NHMB, 2305a).
Draconarius simplicidens Wang, sp. nov.
Figures 59A—B; Map 19
TyprEs.— Female holotype from Pass over Gaoligongshan at 2300 m, Luoshuidong, 28 air km
E TengChong, 24°57’N, 98°45’E, native forest, Baoshan, Yunnan, China (October 26-31, 1998; C.
Griswold, D. Kavanaugh, C-L. Long), deposited in HBI.
ETYMOLOGY.— The specific name refers to the simple epigynum.
DIAGNOSIS.— The female of this new species is similar to D. dubius and D. patellabifidus by
lacking epigynal teeth and having broad spermathecae but can be recognized by the medially situ-
ated spermathecal heads (Figs. 59A—B).
Eye sizes and interdistances: AME 0.10, ALE 0.18, PME 0.18, PLE 0.20; AME-AME 0.05, AME-
ALE 0.05, PME-PME 0.10, PME-PLE 0.11, AME-PME 0.11. Leg measurements: I: 7.70 (2.42,
2.78, 1.60, 0.90); Il: 6.58 (2.02, 2.34, 1.42, 0.80); III: 5.08 (1.58, 1.66, 1.14, 0.70); IV: 7.26 (2.32,
2.60, 1.64, 0.70). Promargin of chelicera with three, retromargin with two to three teeth. Epigynal
teeth absent; atrium small, indistinct; copulatory ducts small, originating posteriorly, extending
mesad of spermathecae; spermathecal heads large, situated mesad of spermathecae; spermathecal
bases broad, widely separated; spermathecal stalks broad, anteriorly diverging (Figs. 59A—B).
WANG: EAST ASIA COELOTINE SPIDERS 549
MaALeE.— Unknown.
DISTRIBUTION.— China (Yunnan) (Map 19).
OTHER MATERIAL EXAMINED.— None.
Draconarius singulatus (Wang et al., 1990)
Figures 60A—E; Map 19
Coelotes singulatus Wang et al., 1990:192, figs. 36-40 (1 male and 2 female types, holotype not indicated,
from Nanshanping, Chenbu, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:378, figs.
224R-S, 227A, 228D.
Draconarius singulatus Wang, 2002:69.
DrAGNosis.— The female of this species is similar to D. gurkha but can be distinguished by
the broad, medially situated copulatory ducts (Figs. 6(0A—B). The male is similar to D. everesti by
the short cymbial furrow and the prolaterally originating embolus but can be distinguished by the
long RTA and the distinct lateral tibial apophysis (Figs. 60C-E).
DESCRIPTION.— Described by Wang et al. (1990). Promargin of chelicera with three teeth,
retromargin with two. Female epigynal teeth short, situated slightly anterior atrial margin; atrium
small; copulatory ducts broad, situated mesad of spermathecae; spermathecal heads not visible
from dorsal view; spermathecae broad, widely separated (Figs. 60A—B). Male palpal patellar
apophysis present, long; RTA slightly longer than half tibial length; lateral tibial apophysis large,
widely separated from RTA; cymbial furrow short; conductor broad, with small basal lamella; con-
ductor dorsal apophysis slender; embolus prolateral in origin; median apophysis elongated, spoon-
like (Figs. 60C-E).
DISTRIBUTION.— China (Hunan) (Map 19).
MATERIAL EXAMINED.— CHINA: Hunan: Chanbu, Nanshanping, July 30, 1982, 1 male and 2
female types (J. F Wang, HBI).
Draconarius stemmleri (Brignoli, 1978)
Figures 61A—B; Map 19
Coelotes stemmleri Brignoli, 1978:43, figs. 15-16 (female holotype from Sha Gogona, Bhutan, in NHMB,
female paratype from Gogona, Kotota, Bhutan, in MCV, examined).
Draconarius stemmleri: Wang, 2002:69.
DIAGNosIs.— The female of this species is similar to D. yadongensis but can be distinguished
by the medially situated spermathecal heads (Figs. 61 A—B).
DESCRIPTION.— Described by Brignoli (1978). Promargin of chelicera with three teeth, retro-
margin with two. Female epigynal teeth short, broad, situated slightly anterior atrial margin; atri-
um small; copulatory ducts small, situated mesad of spermathecae; spermathecal heads large, situ-
ated mesad of spermathecae; spermathecal bases broad, widely separated; spermathecal stalks
broad, strongly expanded anteriorly (Figs. 61 A—B). Male unknown.
DISTRIBUTION.— Bhutan (Map 19).
MATERIAL EXAMINED.— BHUTAN: Sha Gogona, 3100 m, July-August, 1972, female holotype
(NHMB, 2303a). Kotoka, Gogona, 2600-3400 m alt., June 10, 1972, 1 female paratype (MCV).
550 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconarius streptus (Zhu and Wang, 1994), NEw COMBINATION
Map 20
Coelotes streptus Zhu and Wang, 1994:40, figs. 13-14 (female holotype from Kangding, Sichuan, China, in
NBUMS, not examined).— Song, Zhu and Chen, 1999:378, figs. 224V—W.
DIAGNOSIS.— The female of this species is similar to D. syzygiatus in lacking epigynal teeth,
having broad atrium and medially extending spermathecal bases but can be distinguished by the
long, slender, anteriorly converging spermathecal stalks.
DESCRIPTION.— Described by Zhu and Wang (1994). Chelicerae with three promarginal, and
two retromarginal teeth. Female epigynal teeth absent; atrium large; spermathecal heads small, sit-
uated anteriorly; spermathecal bases widely separated, medially extending; spermathecal stalks
slender, laterally extending, anteriorly converging and close together. Male unknown.
DISTRIBUTION.— China (Sichuan) (Map 20).
MATERIAL EXAMINED.— None.
Draconarius striolatus (Wang et al., 1990)
Figure 62A—B; Map 20
Coelotes striolatus Wang et al., 1990:190, figs. 34-35 (female holotype from Yuzhong, Gansu, China, in HBI,
examined).— Song, Zhu and Chen, 1999:378, figs. 225A—B.
Draconarius striolatus: Wang, 2002:69.
DIAGNOsIS.— The female of this species is similar to D. stemmleri but can be distinguished
by the absence of epigynal teeth (Figs. 62A—B).
DESCRIPTION.— Described by Wang et al. (1990). Promargin of chelicera with three teeth,
retromargin with two. Female epigynal teeth absent; atrium large; copulatory ducts small, situated
posteriorly mesad of spermathecae; spermathecal heads long, situated mesad of spermathecae;
spermathecal bases broad, widely separated; spermathecal stalks anteriorly expanded and converg-
ing (Figs. 62A—B). Male unknown.
DISTRIBUTION.— China (Gansu) (Fig. Map 20).
MATERIAL EXAMINED.— CHINA: Gansu: Yuzhong, August 16, 1988, female holotype (J.F.
Wang, HBI).
Draconarius strophadatus (Zhu and Wang, 1991), NEW COMBINATION
Map 20
Coelotes strophadatus Zhu and Wang, 1991:3, figs. 12-13 (female holotype from Huangshan, Anhui, China,
in NBUMS, not examined).— Song, Zhu and Chen, 1999:378, figs. 225C—D.
DIAGNOsIS.— The female of this species is similar to D. huizhunesis by having the long epig-
ynal teeth but can be distinguished by the closely situated spermathecal stalks and widely separat-
ed spermathecal heads.
DESCRIPTION.— See Zhu and Wang (1991). Chelicerae with three promarginal, and two retro-
marginal teeth. Female epigynal teeth situated anteriorly and close together, strongly elongated;
spermathecal bases small, slightly separated; spermathecal stalks close together, convoluted, and
anteriorly diverging. Male unknown.
DISTRIBUTION.— China (Anhui) (Map 20).
MATERIAL EXAMINED.— None.
WANG: EAST ASIA COELOTINE SPIDERS 551
Draconarius subtitanus (Hu, 1992), New Combination
Map 20
Tegenaria pagana: Hu and Li, 1987:283, figs. 20.34 (misidentification).
Coelotes subtitanus Hu, 1992:42, figs. 9-10 (female holotype, 2 female paratypes from Yadong, Tibet, China,
in SDU, not examined).— Hu. 2001:147, figs. 8-57.1-2.
DraAGNosis.— The female is similar to D. himalayaensis and D. altissimus in having the short
epigynal teeth, anteriorly situated spermathecal heads and rounded, closely situated spermathecae
but can be distinguished from D. himalayaensis by the epigynal teeth position (close to atrium),
from D. altissimus by the slightly separated spermathecae.
DESCRIPTION.— See Hu (1992). Chelicerae with three promarginal, and two retromarginal
teeth. Female epigynal teeth short, widely separated, situated near anterior atrium; atrium broad,
posteriorly situated; spermathecal heads situated anteriorly; spermathecal bases widely separated;
spermathecal stalks broad, rounded, slightly separated. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 20).
MATERIAL EXAMINED.— None.
Draconarius syzygiatus (Zhu and Wang, 1994), NEW COMBINATION
Map 21
Coelotes syzygiatus Zhu and Wang, 1994:37, figs. 14 (female holotype and male paratype from Emei Mt.,
Sichuan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:378, figs. 225G—H, 227C, 228F.
DIAGNOsIS.— The female of this species is similar to D. streptus by the absence of epigynal
teeth, the presence of broad atrium, and the medially extending spermathecal bases but can be dis-
tinguished by the strongly spiraled, rounded spermathecal stalks. The male can be easily identified
by the long, posteriorly extending conductor.
DESCRIPTION.— See Zhu and Wang (1994). Chelicerae with three promarginal, and two retro-
marginal teeth. Female epigynal teeth absent; atrium large; spermathecal heads small, situated ante-
riorly; spermathecal bases widely separated, medially extending; spermathecal stalks long, spi-
raled, rounded. Male palpal patellar apophysis small; RTA short; lateral tibial apophysis long, slen-
der; cymbial furrow about half cymbial length; conductor strongly elongate (about the cymbial
length), posteriorly extended and looped, with broad base and slender apex; conductor lamella
small; conductor dorsal apophysis present; embolic base small; embolus long, slender, strongly
extended posteriorly and then curved back anteriorly; median apophysis small, spoon-like.
DISTRIBUTION.— China (Sichuan)(Map 21).
MATERIAL EXAMINED.— None.
Draconarius terebratus (Peng and Wang, 1997), NEW COMBINATION
Figures 63A—E; 96G-H; Map 21
Coelotes terebratus Peng and Wang, 1997:330, figs. 27-31. (female holotype and male allotype from
Tianpingshan, Sangzhi, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:378, figs.
225M-N, 227E, 228H.
DIAGNOsISs.— The female of this species is similar to D. ornatus by lacking epigynal teeth,
having posteriorly situated epigynal hoods, anteriorly extending copulatory ducts, and long sper-
mathecal heads but can be distinguished by the rounded atrium and the broad spermathecal bases
552 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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(Figs. 63A—B). The male can be recognized by the large lateral tibial apophysis, the lobed embol-
ic base, and the long, toothed embolus (Figs. 63C-E).
DESCRIPTION.— Described by Peng and Wang (1997). Promargin of chelicera with three teeth,
retromargin with two. Female lacking epigynal teeth; atrium large; epigynal hoods situated poste-
riorly laterad of atrium; copulatory ducts posteriorly originating, strongly expanded anteriorly;
spermathecal heads long, slender, anteriorly extending; spermathecal bases small, widely separat-
ed; spermathecal stalks anteriorly diverging (Figs. 63A—B). Male palpal patellar apophysis present;
RTA long; lateral tibial apophysis large, widely separated from RTA; cymbial furrow short; con-
ductor long, broad, with slender, posteriorly hooked apex and small basal lamella; conductor dor-
sal apophysis small; embolic base lobed; embolus posterior in origin, modified with a small tooth;
median apophysis small, spoon-like (Figs. 63C—E).
DISTRIBUTION.— China (Hunan) (Map 21).
MATERIAL EXAMINED.— CHINA: Hunan: Sangzhi, Tianpingshan, October 16, 1986, female
holotype and male allotype (J.F. Wang, HBI).
Draconarius tibetensis Wang, sp. nov.
Figures 64A—C; Map 21
TyprE.— Male holotype from Yupik Valley, 29.48°N, 96.14°E, Tibet, China (May 14, 1998; G
Schaller), deposited in AMNH.
ETYMOLOGyY.— The specific name refers to the type locality.
DIAGNOsIS.— The male of this species is similar to D. baronii but can the distinguished by the
broad dorsal edge of conductor (Figs. 64A—C).
MALE.— Total length 11.2. Carapace 6.80 long, 3.40 wide. Abdomen 4.40 long, 3.20 wide.
Eye sizes and interdistances: AME 0.14, ALE 0.20, PME 0.15, PLE 0.17; AME-AME 0.14, AME-
ALE 0.14, PME-PME 0.28, PME-PLE 0.30, AME-PME 0.18. Leg measurements: I: 16.8 (4.20,
5.60, 4.40, 2.60); II: 15.6 (4.00, 5.60, 4.20, 2.40); II: 14.2 (3.60, 4.40, 4.00, 2.20); 1V: — (4.40, —
, —, —). Chelicerae with three promarginal, and two widely separated retromarginal teeth. Male
palp with large patellar apophysis; RTA slightly longer than half tibial length; lateral tibial apoph-
ysis widely separated from RTA; cymbial furrow short; conductor short, with broad dorsal edge;
conductor dorsal apophysis small; conductor lamella large; embolus posterior in origin; median
apophysis spoon-like, elongated (Figs. 64A—C).
FEMALE.— Unknown.
DISTRIBUTION.— China (Tibet) (Map 21).
OTHER MATERIAL EXAMINED.— None.
Draconarius trifasciatus (Wang and Zhu, 1991)
Map 22
Coelotes trifasciatus Wang and Zhu, 1991:3, figs.1—4 (female holotype and male paratype from Mt. Emei,
Sichuan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:388, figs. 225U—V, 2271, 228J.
Draconarius trifasciatus: Wang, 2002:69.
DIAGNOsIS.— The female of this species is similar to D. stemmleri but can be distinguished
by the less distinct spermathecal heads and the anteriorly converging spermathecae. The male can
be recognized by the large cymbial furrow, the long conductor, and the toothed (lobed) embolic
base.
WANG: EAST ASIA COELOTINE SPIDERS 553
DESCRIPTION.— See Wang and Zhu (1991). Chelicerae with three promarginal, and two retro-
marginal teeth. Female epigynum with widely separated, posteriorly situated, broad epigynal teeth;
atrium broad, situated posteriorly near epgastric furrow; spermathecae broad. Male palp with patel-
lar apophysis; RTA long; lateral tibial apophysis situated near RTA; cymbial furrow large, almost
as long as cymbium: conductor long, slender; conductor dorsal apophysis present; embolic base
with a prolateral lobe; embolus long, posterior in origin.
DISTRIBUTION.— China (Sichuan) (Map 22).
MATERIAL EXAMINED.— None.
Draconarius tryblionatus (Wang and Zhu, 1991), NEw COMBINATION
Map 21
Coelotes tryblionatus Wang and Zhu, 1991:3, figs. 5-8 (female holotype, male and female paratypes from Mt.
Qingcheng, Sichuan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:388, figs. 226C—D,
DIK 2281.
DIAGNOsIS.— The female of this species is similar to D. streptus and D. syzygiatus in lacking
epigynal teeth and having broad atrium but can be distinguished by the laterally extending sper-
mathecal bases. The male is similar to D. uncinatus in having the bifurcate patellar apophysis, a
broad conductor, and a strong embolus but can be distinguished by the spiraled embolus and the
broad median apophysis.
DESCRIPTION.— See Wang and Zhu (1991). Chelicerae with three promarginal, and three
retromarginal teeth. Female without epigynal teeth; atrium large; spermathecal bases widely sepa-
rated, laterally extending; spermathecal stalks laterally extending and then curved medially, ante-
riorly converging and close together. Male palp with bifurcate patellar apophysis; RTA long; later-
al tibial apophysis present; cymbial furrow short; conductor broad; median apophysis situated near
conductor; embolus strong, spiraled.
DISTRIBUTION.— China (Sichuan) (Map 21).
MATERIAL EXAMINED.— None.
Draconarius uncinatus (Wang et al., 1990), NEW COMBINATION
Figures 65A—C; 96F; Map 22
Coelotes uncinatus Wang et al., 1990:188, figs. 29-31 (2 male types, holotype not indicated, from Tianmu-
shan, Zhejiang, China, in HBI, examined).— Song, Zhu and Chen, 1999:388, figs. 227L, 228M.
DiAGNosIs.— This species is similar to D. tryblionatus in having a bifurcate patellar apoph-
ysis, a broad conductor, and a strong embolus but can be distinguished by the non-spiraled embo-
lus and the small median apophysis (Fig. 65A—C).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Male palp with bifurcate patellar apophysis; RTA long; lateral tibial apophysis
present; cymbial furrow short; conductor broad; conductor dorsal apophysis small; conductor
lamella small; embolus posterior in origin, broad; median apophysis spoon-like, small (Figs.
65A—C). Female unknown.
DISTRIBUTION.— China (Zhejiang) (Map 22).
MATERIAL EXAMINED.— CHINA: Zhejiang: Tianmushan, October 15, 1974, 2 male types (J.F.
Wang, HBI).
554 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconarius venustus Ovtchinnikov, 1999
Map 22
Draconarius venustus Ovtchinnikov, 1999:70, figs. 23-27 (male holotype and female paratype from
Khszratisho Mt., Yachsuriver Valley, Tajikistan, deposited in cSO, not examined).— Wang, 2000:69.
DIAGNOsIS.— Similar to D. wudangensis in having the medially situated and widely separat-
ed epigynal teeth, and similar spermathecal tubes but can be distinguished by the posteriorly situ-
ated spermathecal heads and the strong anterior expansion of spermathecae of female, and by the
long RTA (almost tibial length), short tibia (about patellar length), and the short patellar apophysis
of male.
DESCRIPTION.— See Ovtchinnikov (1999). Chelicerae with three promarginal, and two retro-
marginal teeth. Female epigynal teeth short, widely separated; atrium small, situated posteriorly
near epigastric furrow; copulatory ducts apparent, originating posteriorly mesad of spermathecae;
spermathecal heads situated posteriorly, near spermathecal bases; spermathecal bases widely sep-
arated; spermathecal stalks strongly converging and expanded anteriorly. Male palp with patellar
apophysis small; RTA almost as long as tibia; lateral tibial apophysis present; cymbial furrow
almost as long as cymbium; conductor short, with large lamella; conductor dorsal apophysis pres-
ent; embolus posterior in origin, long; median apophysis spoon-like.
DISTRIBUTION.— Tajikistan (Map 22).
MATERIAL EXAMINED.— None.
Draconarius wenzhouensis (Chen, 1984)
Figures 66A—B; Map 22
Coelotes wenzhouensis Chen, 1984:3, figs. 7-8 (female holotype and | female paratype from Xueshan,
Wenzhou, Zhejiang, China, in HTC, examined).— Chen and Zhang, 1991:190, figs. 189.1—-2;— Song, Zhu
and Chen, 1999:388, figs. 226I-J.
Draconarius wenzhouensis: Wang, 2002:69.
DIAGNOSIS.— This species is similar to D. labiatus but can be distinguished by the large sper-
mathecal heads and the anteriorly extending spermathecae (Fig. 66A—B).
DESCRIPTION.— Described by Chen (1984). Cheliceral promargin with three teeth, retromar-
gin with two. Female epigynal teeth short, close together, anteriorly situated; atrium situated pos-
teriorly near epigastric furrow, with anterior atrial margin lip-shaped and expanded posteriorly;
copulatory ducts posteriorly originating, extending mesad of spermathecae; spermathecal heads
large; spermathecal bases widely separated; spermathecal stalks strongly expanded and converging
anteriorly (Figs. 66A—B). Male unknown.
DISTRIBUTION.— China (Zhejiang) (Map 22).
MATERIAL EXAMINED.— CHINA: Zhejiang: Wenzhou, Xueshan, March 10—12, 1980, female
holotype and 2 female paratypes (Z.F. Chen, HTC).
Draconarius wudangensis (Chen and Zhao, 1997)
Figures 67A—E; Map 23
Coelotes wudangensis Chen and Zhao, 1997:87, figs. |—4 (1 male and | female paratypes, holotype not indi-
cated, from Jinding, Wudangshan, Hubei, China, in HUW, examined).— Song, Zhu and Chen, 1999:388,
figs. 226K-L, 2270, 229A.
Draconarius wudangensis: Wang, 2002:69.
Draconarius parawudangensis Zhang, Zhu and Song, 2002:53, figs. 5— 6. NEW SYNONYMY.
WANG: EAST ASIA COELOTINE SPIDERS 555
DIAGNosIs.— The female is similar to D. aspinatus, D. calcariformis, D. coreanus, D. davidi,
D. linzhiensis, D. picta, D. gingzangensis, and D. venustus in having similar spermathecae but can
be distinguished by the medially situated, widely separated (widely separated from atrium) epigy-
nal teeth, the anteriorly situated (anterior 1/3 of spermathecae) spermathecal heads, and the broad
spermathecal bases (Figs. 67A—B). The male is similar to D. venustus but can be distinguished by
the short RTA (half tibial length), long tibia (twice patellar length), and the long patellar apophysis
(Figs. 67C-E).
DESCRIPTION.— Described by Chen and Zhao (1997) and Wang (2002). Cheliceral promargin
with three teeth, retromargin with two. Female epigynal teeth short, widely separated; atrium small,
situated posteriorly near epigastric furrow; copulatory ducts originating posteriorly, extending
mesad of spermathecae; spermathecal heads situated anteriorly; spermathecal bases widely sepa-
rated, broad; spermathecal stalks broad, anteriorly extending and converging (Figs. 67A—B). Male
palp with patellar apophysis long; RTA approximately half tibial length; lateral tibial apophysis
present; cymbial furrow long, slightly more than half cymbial length; conductor short, with large
basal lamella; conductor dorsal apophysis present; embolus posterior in origin, long; median
apophysis spoon-like, elongated (Figs. 67C-E).
DISTRIBUTION.— China (Hubei, Shaanxi, Shanxi) (Map 23).
MATERIAL EXAMINED.— CHINA: Hubei: Wudangshan, Jinding, August 1996, 1 male and 1
female paratypes (J. Chen, HUW); Wudangshan, Jinding, September 24, 1997, 7 females (X.P.
Wang, IZB); Wudangshan, Nanya to Jinding, September 24, 1997, | male and 16 females (X.P.
Wang, IZB). Shaanxi: Taibaishan, Haoping, August 11, 1989, 1 male and | female, 1 male and 1
female /1 male and | female, 1 male (X.P. Wang, AMNH and MCB); Taibaishan, Mingxinshi, Aug.
8, 1989, 1 female (X.P. Wang, IZB). Shanxi: Yongji, July 20, 1980, 1 female (M.S. Zhu, HUB, No-
044).
Draconarius wuermlii (Brignoli, 1978)
Figures 68A—B; Map 23
Coelotes wuermlii Brignoli, 1978:44, figs. 21-22 (female holotype from Dechhi Paka, Bhutan, in NHMB,
examined).
Paracoelotes wuermlii: Brignoli, 1982:349.
Draconarius wuermlii: Wang, 2002:69.
DIAGNOsIs.— The female of this species is similar to D. pseudowuermlii but can be distin-
guished by the looped copulatory ducts, the less convoluted spermathecae, and the small spermath-
ecal bases (Figs. 68A—B).
DESCRIPTION.— Described by Brignoli (1978). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth short, widely separated, situated posteriorly and laterad of
atrium; atrium small; copulatory ducts originating posteriorly and mesad of spermathecae, looped
around spermathecae; spermathecal heads situated anteriorly; spermathecal bases widely separat-
ed; spermathecal stalks long, anteriorly converging (Figs. 68A—B). Male unknown.
DISTRIBUTION.— Bhutan (Map 23).
MATERIAL EXAMINED.— BHUTAN: Dechhi Paka, 3300 m, June 20, 1972, female holotype
(Basel-Bhutan Expediture 1972, NHMB, 2304a).
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Draconarius yadongensis (Hu and Li, 1987), NEW COMBINATION
Map 23
Wadotes yadongensis Hu and Li, 1987:280, figs. 20.1—2 (female holotype from Yadong, Tibet, China, in SDU,
not examined).— Song, Zhu and Chen, 1999:395, figs. 230M-N:— Hu, 2001:153, figs. 8-61.1-2.
Coelotes yadongensis: Hu, 1992:43.
DIAGNOSIS.— The female of this species is similar to D. stemmleri but can be distinguished
by the anteriorly situated spermathecal heads.
DESCRIPTION.— See Hu and Li (1987). Chelicerae with three promarginal, and two retromar-
ginal teeth. Female epigynal teeth short, broad, situated slightly anterior of atrial margin; atrium
small; spermathecal bases broad, widely separated; spermathecal stalks broad, strongly expanded
anteriorly. Male unknown.
DISTRIBUTION.— China (Tibet) (Map 23).
MATERIAL EXAMINED.— None.
Draconarius yichengensis Wang, sp. nov.
Figures 69A—B; Map 23
Types.— Female holotype from Yicheng, Shanxi, China (August 15, 1983; M.S. Zhu),
deposited in HUB (No-83-—0025).
ETYMOLOGY.— The specific name refers to the type locality.
DIAGNOSIS.— The female of this new species can be easily recognized by the long epigynal
teeth, the broad, rounded spermathecae, and the ventrally originating spermathecal heads (Figs.
69A-B).
FEMALE.— Total length 6.20. Carapace 3.12 long, 2.05 wide. Abdomen 3.08 long, 1.81 wide.
Eye sizes and interdistances: AME 0.08, ALE 0.15, PME 0.11, PLE 0.12; AME-AME 0.09, AME-
ALE 0.15, PME-PME 0.13, PME-PLE 0.16. Leg measurements: I: 7.94 (2.24, 2.81, 1.72, 1.17); U:
7A3 (2.10, 2.54; 1.65, 1.14); IM: 7.06 (1.95, 2.32, 1.73, 0:94); IV: 9:57 (2.56; 3311 2020):
Promargin of chelicera with three teeth, retromargin with two. Female epigynal teeth long, situat-
ed near anterior atrial margin; atrium broad; copulatory ducts small; spermathecal heads anterior-
ly situated, originating ventrally; spermathecal bases broad, widely separated; spermathecal stalks
broad, rounded, anteriorly converging (Figs. 69A—B). Male unknown.
MALE.— Unknown.
DISTRIBUTION.— China (Shanxi) (Map 23).
OTHER MATERIAL EXAMINED.— None.
Draconarius yostianus (Nishikawa, 1999)
Map 24
Coelotes yosiianus Nishikawa, 1999:23, figs. 1-5 (female holotype and 1 female paratype from Jiabao Dong,
Xingren Cun, Jiazhuan Xiang, Bama Xian, Guangxi, China, in NSMT, not examined).
Draconarius yosiianus: Wang, 2002:69.
DIAGNOsIS.— The female of this species can be easily recognized by the absence of eyes, the
widely separated, medially expanded spermathecae.
DESCRIPTION.— See Nishikawa (1999). Chelicerae with three promarginal, and two retromar-
ginal teeth. Female epigynal teeth short, situated slightly anterior atrium; atrium large; spermathe-
cal bases widely separated; spermathecal stalks widely separated, medially expanded. Male
unknown.
WANG: EAST ASIA COELOTINE SPIDERS 557
DISTRIBUTION.— China (Guangxi) (Map 24). Known only from a cave.
MATERIAL EXAMINED.— None.
Genus Femoracoelotes Wang, 2002
Femoracoelotes Wang, 2002:81 (type species, by original designation, Coelotes platnicki Wang and Ono,
1998 from China).
DIAGNOsIS.— The female of this genus can be easily recognized by the absence of epigynal
teeth and the presence of broad copulatory ducts, the male by the presence of a femoral apophysis
and the absence of median apophysis (Figs. 70-71).
PHYLOGENETIC PLACEMENT.— The presence of four cheliceral retromargin teeth suggests
Femoracoelotes is the sister group of Coronilla from China, together they form the sister group of
all other coelotines (Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China (Map 25).
COMPOSITION.— 2 species:
1. Femoracoelotes latus (Wang, Tso and Wu, 2001)
2. Femoracoelotes platnicki (Wang and Ono, 1998)
Femoracoelotes latus (Wang, Tso and Wu, 2001)
Figures 7OA—E; Map 25
Coelotes latus Wang, Tso and Wu, 2001:130, figs. 11-21 (male holotype, 1 male and 1 female paratypes from
Nantou, Taiwan, in THU, examined).
Femoracoelotes latus: Wang, 2002:81.
DIAGNOsIs.— This species can be distinguished from F: platnicki by the small, slightly
expanded copulatory ducts, the less convoluted spermathecae of female (Figs. 70A—B), and by the
short conductor and the short, and dorsally situated conductor dorsal apophysis (Figs. 70C-E).
DESCRIPTION.— Described by Wang, Tso and Wu (2001). Chelicerae with three promarginal
and four retromarginal teeth. Epigynal teeth absent; atria small; copulatory ducts broad, slightly
extending anteriorly; spermathecal heads small, anteriorly situated; spermathecal bases broad,
widely separated; spermathecal stalks slightly elongated, not convoluted (Figs. 70A—B). Male palp
with femoral apophysis slightly bifurcate; femora with numerous short spines on distal prolateral
surface; patellar apophysis short; RTA as long as tibia; lateral tibial apophysis absent; cymbial fur-
row short; conductor short, broad, with small basal lamella; conductor dorsal apophysis short;
median apophysis absent; embolus strong, prolateral to posterior in origin, with bifurcate apex
(Figs. 7OC-E).
DISTRIBUTION.— China (Taiwan) (Map 25).
MATERIAL EXAMINED.— CHINA: Taiwan: Nantou, Hui-Sun Experimental Forest Station,
October 1997, male holotype, 1 male and | female paratypes (Hai- Ying Wu, THU).
Femoracoelotes platnicki (Wang and Ono, 1998)
Figures 71A—E; Map 25
Coelotes platnicki Wang and Ono, 1998:148, figs. 15-19 (male holotype and female paratype from Mt.
Tengchih, Paoshan-tsun, Taoyuan-hsiang, Kaohsiung-hsien, Taiwan, in NSMT, examined).— Song, Zhu
and Chen, 1999:377.
Femoracoelotes platnicki: Wang, 2002:82, figs. 227-241.
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DIAGNOSIS.— This species can be distinguished from F. latus by the broad, anteriorly expand-
ed copulatory ducts, the convoluted spermathecae of female (Figs. 71A—B), and by the broad,
strongly elongated conductor and the long, ventrally situated conductor dorsal apophysis of male
(Figs. 71C-E).
DESCRIPTION.— Described by Wang and Ono (1998). Chelicerae with three promarginal and
four retromarginal teeth (occasionally five). Female epigynal teeth absent; atrium small, situated
posteriorly near epigastric furrow: copulatory ducts broad, anteriorly expanded; spermathecal
heads large; spermathecal bases widely separated; spermathecal stalks short, convoluted (Figs.
71A-B). Male palp with femoral apophysis bifurcate; femora with numerous short spines on dis-
tal prolateral surface; patellar apophysis present: RTA as long as tibia; lateral tibial apophysis
absent; cymbial furrow short; conductor long, broad, strongly modified, lamella small, dorsal
apophysis long, ventrally situated; median apophysis absent; embolus strong, prolateral to posteri-
or in origin, with slightly modified apex (Figs. 71C—E).
DISTRIBUTION.— China (Taiwan) (Map 25).
MATERIAL EXAMINED.— CHINA: Taiwan: Kaohsiung-hsien, Taoyuan-hsiang, Paoshan-tsun,
Mt. Tengchih, 1550-1800 m alt., November 1, 1989, 5 males and 7 females (H. Ono, NSMT,
NSMT-Ar.3421); Kaohsiung-hsien, Taoyuan-hsiang, Paoshan-tsun, Mt. Tengchih, 1550 m alt.,
November 1, 1989, 1 male (H. Ono, NSMT, NSMT-Ar.3430); Kaohsiung-hsien, Taoyuan-hsiang,
Paoshan-tsun, Mt. Tengchih, 1550-1800 m alt.. November 1, 1989, male holotype and female
paratype (H. Ono, NSMT, NSMT-Ar.3421).
Genus Leptocoelotes Wang, 2002
Leptocoelotes Wang, 2002:105 (type species, by original designation, Coelotes pseudoluniformis Zhang, Peng
and Kim, 1997 from China).
DIAGNOsIs.— The female can be easily recognized by the broad, weakly sclerotized epigynal
teeth, the shallow atrium, and the short copulatory ducts, and the male by the complex conductor,
the absence of a conductor dorsal apophysis, and the absence of a median apophysis (Figs. 72-73).
PHYLOGENETIC PLACEMENT.— Remains unresolved with Tegecoelotes and the clade with
spoon-like median apophysis and slender epigynal teeth (Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China (Map. 26).
COMPOSITION.— 2 species:
1. Leptocoelotes edentulus (Wang and Ono, 1998)
2. Leptocoelotes pseudoluniformis (Zhang, Peng and Kim, 1998)
Leptocoelotes edentulus (Wang and Ono. 1998)
Figures 72A—B; Map 26
Coelotes edentulus Wang and Ono, 1998:142, figs. 1-2 (female holotype from Ilan, Taiwan, in NSMT, exam-
ined).— Song, Zhu and Chen, 1999:375.
Leptocoelotes edentulus: Wang, 2002:105.
DIAGNOSIS.— The female of this species can be distinguished from L. pseudoluniformis by the
anteriorly situated epigynal hoods and the posteriorly originating copulatory ducts (Figs. 72A—B).
DESCRIPTION.— Described by Wang and Ono (1998). Chelicerae with five to six promarginal
and five retromarginal teeth. Female epigynal teeth broad but weak, indistinct; atrium broad; epig-
ynal hoods deep, situated anterad of atrium; copulatory ducts short, broad, originating posteriorly
WANG: EAST ASIA COELOTINE SPIDERS 559
and laterad of spermathecae; spermathecal heads small, situated anteriorly, laterally extending;
spermathecal bases broad, slightly separated; spermathecal stalks short, close together (Figs.
72A—B). Male unknown.
DISTRIBUTION.— China (Taiwan) (Map 26).
MATERIAL EXAMINED.— China: Taiwan: Ilan, Tienking-miao, near Tali, 50 m, March 17,
1991, female holotype (H. Ono, NSMT, NSMT-Ar.3427); Nantou, Tatachia, 2100 m, March 5,
1991 1 female paratype (H. Ono, NSMT, NSMT-Ar.3427).
Leptocoelotes pseudoluniformis (Zhang, Peng and Kim, 1997)
Figures 73A—G; Map 26
Coelotes pseudoluniformis Zhang, Peng and Kim, 1997:293, fig. 6-7 (female holotype from Tiantong,
Zhejiang, China, in HBI, examined).
Leptocoelotes pseudoluniformis: Wang, 2002:105.
DIAGNOsIS.— The female of this species can be recognized from L. edentulus by the laterally
situated epigynal hoods and the anteriorly originating copulatory ducts (Figs. 73A—D). The male
can be distinguished by the tiny lateral tibial apophysis, the strongly bifurcate conductor, the
absence of conductor dorsal apophysis, and the absence of median apophysis (Figs. 73E-G).
DESCRIPTION.— Described by Zhang, Peng and Kim (1997). Chelicerae with five promargin-
al and four to five retromarginal teeth. Female epigynal teeth broad but weak, situated on anterior
atrial margin; atrium broad; epigynal hoods deep, situated laterad of atrium; copulatory ducts short,
originating anteriorly and laterad of spermathecae; spermathecal heads small, situated anteriorly;
spermathecal bases broad, widely separated; spermathecal stalks short, broad, widely separated
(Figs. 73A—D). Male palp with patellar apophysis small, dorsally curved; RTA approximately half
tibial length; lateral tibial apophysis tiny, situated posteriorly near RTA; cymbial furrow short; con-
ductor strongly bifurcate, slightly spiraled; conductor dorsal apophysis absent, lamella small;
embolus posterior in origin; median apophysis absent (Figs. 73E-G).
DISTRIBUTION.— China (Zhejiang, Hunan) (Map 26).
MATERIAL EXAMINED.— CHINA: Zhejiang: Tiantong, January 20, 1988, female holotype (Y.J.
Zhang, HBI). Hunan: Changsha, Yuelushan, Dec. 22, 1982, 2 males and 4 females (J.F. Wang,
HBI).
Genus Longicoelotes Wang, 2002
Longicoelotes Wang, 2002:109 (type species, by original designation, Longicoelotes karschi Wang, 2002).
DiAGnosis.— The female can be easily recognized by the absence of epigynal teeth and the
distinct shape of epigynum, and the male by the strongly elongated patellar apophysis and the
reduced median apophysis (Figs. 74A—E).
PHYLOGENETIC PLACEMENT.— The reduced atrium and the presence of small copulatory ducts
support the sister group relationship between Longicoelotes and all coelotines with epigynal teeth
(Wang 2002).
DESCRIPTION.— See Wang (2002).
DIsTRIBUTION.— China, Senkaku (Map. 27).
COMPOSITION.— 3 species, including 2 new combinations:
1. Longicoelotes karschi (Wang, 2002)
2. Longicoelotes kulianganus (Chamberlin, 1924), NEw COMBINATION
3. Longicoelotes senkakuensis (Shimojana, 2000), NEw COMBINATION
560 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Longicoelotes karschi Wang, 2002
Figures 74A—E; Map 27
Coelotes mollendorffi: Schenkel, 1963:280, fig. 158.— Chen and Zhang, 1991:187, fig. 185 (misidentifica-
tion).
Coelotes moellendorffi: Song, Zhu and Chen, 1999:376, figs. 221E—-F, 222M, 224A (misidentification).
Longicoelotes karschi Wang, 2002:109.
DIAGNOSIS.— Same as for genus (Figs. 74C-E).
DESCRIPTION.— Described by Wang (2002). Chelicerae with three promarginal, two retromar-
ginal teeth. Female without epigynal teeth; middle epigynum relatively elevated with clear lateral
edges which converge anteriorly; atrium small; copulatory ducts short, originating and situated lat-
erad of spermathecae; spermathecal heads apparent, extending laterally; spermathecal bases broad,
slightly separated; spermathecal stalks short, situated close together (Figs. 74A—B). Male palp with
patellar apophysis strongly elongated, longer than tibial length; RTA almost as long as tibia; later-
al tibial apophysis widely separated from RTA, situated relatively dorsally; cymbial furrow short;
conductor short, broad, with a dorsally bifurcate apophysis and ventrally broad apophysis; conduc-
tor lamella small; conductor dorsal apophysis present; median apophysis reduced to a small apoph-
ysis, not spoonlike; embolus basal in origin (Figs. 74C—E).
DISTRIBUTION.— China (Jiangsu, Zhejiang) (Map 27).
MATERIAL EXAMINED.— CHINA: Zhejiang: West Tschenkiang, April 1872, 1 female (A.
David, MNHN, B2011 bis); Hangtscheou, 1925, 3 females (MNHN, B2011 bis); Lin-An, October
19, 1974, 3 males and 3 females (C.D. Zhu, NBUMS). Jiangsu: Nanjin, Zijin (Purple) Mt., 350—
450 m, September 13, 1997, male holotype (X.P. Wang, AMNH); Nanjin, Zijin (Purple) Mt.,
October 9, 1988, | female paratype (P. Beron, AMNH).
Longicoelotes kulianganus (Chamberlin, 1924), NEW COMBINATION
Map 27
Coelotes kulianganus Chamberlin, 1924:24, fig. 40 (female holotype from Kuliang, Fujian, China, in USNM,
examined, with the abdomen missing).
DIAGNOsISs.— The female epigynum of this species is similar to L. karschi (according to
Chamberlin, 1924). Unfortunately, the holotype lacks the abdomen. Further collection of both male
and female from the type locality is needed.
DESCRIPTION.— See Chamberlin (1924). Female chelicerae with three promarginal, two retro-
marginal teeth. Male unknown.
DISTRIBUTION.— China (Fujian) (Map 27).
MATERIAL EXAMINED.— CHINA: Fujian: Kuliang, 2400 m, female holotype (with abdomen
missing) (N. Gist Gee, USNM, No. 883).
Longicoelotes senkakuensis (Shimojana, 2000), NEW COMBINATION
Map 27
Coelotes senkakuensis Shimojana, 2000:175, figs. 44-49 (female holotype from Senkaku, deposited in
NSMT, not examined).
DIAGNOsIS.— The female of this species appears similar to L. karschi on the basis of the pub-
lished description but can be distinguished by the small spermathecal heads and the differences in
shapes between their epigynum.
WANG: EAST ASIA COELOTINE SPIDERS 561
DESCRIPTION.— See Shimojana (2000). Female chelicerae with three promarginal, two retro-
marginal teeth. Male unknown.
DISTRIBUTION.— Senkaku (Map 27).
MATERIAL EXAMINED.— None.
Genus Platocoelotes Wang, 2002
Platocoelotes Wang, 2002:119 (type species, by original designation, Coelotes impletus Peng and Wang, 1997
from China).
DIAGNOsIS.— The female can be easily recognized by the absence of epigynal teeth, the pres-
ence of an anteriorly situated epigynal cavity (depression), the deep, posteriorly situated epigynal
hoods, and the long, strongly convoluted spermathecae, and the male by the presence of two patel-
lar apophyses, the elongated cymbial furrow, the presence of a conductor posterior apophysis, the
long embolus, and the absence of median apophysis (Figs. 75-78).
PHYLOGENETIC PLACEMENT.— The absence of both epigynal teeth and median apophysis sup-
ports the sister group relationship between Platocoelotes and Spiricoelotes, together they remain
unresolved with the genera Draconarius, Asiacoelotes, and Bifidocoelotes (Wang, 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China (Map 28).
COMPOSITION.— 5 species, including | new species:
1. Platocoelotes impletus (Peng and Wang, 1997)
2. Platocoelotes icohamatoides (Peng and Wang, 1997)
3. Platocoelotes icohamatus (Zhu and Wang, 1991)
4. Platocoelotes kailiensis Wang, sp. nov.
. Platocoelotes lichuanensis (Chen and Zhao, 1998)
Nn
Key to the Species of the Genus Platocoelotes
1. Males (those of P. icohamatus and P. icohamatoides unknown)...................00000: py
Females (those of P. lichuanensis unknown, and P. icohamatus not examined)............ 4
Pebmbolus with basevextending prolaterally (Figss 7/527 SA). es ae ce ee ee 3)
Emibolusswithibaseextendins postenorly(Fisav/ Du nana s a ee eee kailiensis
se Conductor with apical apophyses large, as in’ Fig. J8A..... 2.5... s ee ee lichuanensis
Conductor with-apicallapophyses small asim Fig. JOD) a2 eicce ie os cusses aes impletus
4. Spermathecal bases large, transversely extending; spermathecal stalks broad, with less than
EMECE SLOOP Ss (RIGA) seis he Set et AE ePID S02 le a Ut RO EE Sted aaa impletus
Spermathecal bases small, close together, longitudinally extending, spermathecal stalks narrow,
Wathratleasethree loops (Eas; 7OB< 07 Bis asicioratmura cretion cis eine Sew es 5
5. Spermathecal stalks extremely long, with at least five loops (Fig. 77B)........... kailiensis
Spermathecal stalks moderately long, with 3-4 loops (Fig. 76B)............ icohamatoides
Platocoelotes impletus (Peng and Wang, 1997)
Figures 75A—-E; Map 28
Coelotes impletus Peng and Wang, 1997:328, figs. 11-19 (2 male and 2 female types from Zhangjiajie, Hunan,
China, in HBI, examined).— Song, Zhu and Chen, 1999:375, figs. 220D-E, 222H, 223K.
Platocoelotes impletus: Wang, 2002:122, figs. 336-348.
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DIAGNOsSIS.— The female of this species is similar to P. icohamatus, unfortunately the vulva
of P. icohamatus was not illustrated in the original paper and the specimens of P. icohamatus were
not available for this research. The female can be distinguished from other Platocoelotes by the
anteriorly diverging spermathecae and less looped copulatory ducts (two loops) (Figs. 75 A—B). The
male can be differentiated from P. kailiensis by the prolaterally extending embolic base, and from
P. lichuanensis only by the smaller conductor apical apophyses (Figs. 75C—E).
DESCRIPTION.— See Peng and Wang (1997) and Wang (2002). Chelicerae with three promar-
ginal and two retromarginal teeth. Female epigynum without teeth; epigynal hoods situated poste-
riorly, near epigastric margin; atrium large, longitudinally elongated; epigynum with anteriorly sit-
uated cavity; copulatory ducts short; spermathecal heads small; spermathecal bases extending
transversely; spermathecal stalks broad, with less than three loops (Figs. 75A—B). Male palp with
two patellar apophyses; retrolateral tibial apophysis long, occupying almost entire tibial length; lat-
eral tibial apophysis small, widely separated from RTA; cymbial furrow about half cymbial length
or slightly shorter; conductor broad, with a long, posteriorly extending apophysis and an anterior-
ly extending finger-like apophysis; conductor dorsal apophysis broad; conductor basal lamella
small; embolus posterior in origin, long; median apophysis absent (Figs. 75C—E).
DISTRIBUTION.— China (Hunan) (Map 28).
MATERIAL EXAMINED.— CHINA: Hunan: Zhangyjiajie, August 18-20, 198?, 2 male and 2
female types (J.F. Wang and X.J. Peng, HBI).
Platocoelotes icohamatoides (Peng and Wang, 1997)
Figures 76A—B; Map 28
Coelotes icohamatoides Peng and Wang, 1997:328, figs. 5-10 (1 female paratype from Naer Mt., Fenghuang,
Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:375, figs. 219Q-R.
Platocoelotes icohamatoides: Wang, 2002:122.
DIAGNOsIS.— The female of this species is similar to P. kailiensis but can be distinguished by
the less looped spermathecal stalks (with 3 loops) (Figs. 76A—B).
DESCRIPTION.— Described by Peng and Wang (1997). Chelicerae with three promarginal and
two retromarginal teeth. Female epigynum without epigynal teeth; epigynal hoods situated poste-
riorly near epigastric margin; atrium large, longitudinally elongated; epigynum with anteriorly sit-
uated cave; copulatory ducts short; spermathecal heads small; spermathecal bases extending longi-
tudinally, situated close together; spermathecal stalks broad, with at least three loops (Figs.
76A—B). Male unknown.
DISTRIBUTION.— China (Hunan) (Map 28).
MATERIAL EXAMINED.— CHINA: Hunan: Fenghuang, Naer Mt., | female paratype (J.F. Wang,
HBI).
Platocoelotes icohamatus (Zhu and Wang, 1991)
Map 28
Coelotes icohamatus Zhu and Wang, 1991:2, fig. 5-7 (female holotype, male and female paratypes from
Sichuan, China, in NBUMS, not examined).— Song, Zhu and Chen, 1999:375, figs. 220A, 222F, 2231.
Platocoelotes icohamatus: Wang, 2002:122.
DIAGNOsIS.— The female epigynum of this species is identical to P. impletus, unfortunately
the vulva was not illustrated in the original paper and the specimens were not available for this
research. The male paratypes are similar to D. calcariformis and might be mistakenly matched.
WANG: EAST ASIA COELOTINE SPIDERS 563
Nortes.— Judging from the illustrations by Zhu and Wang (1991, figures 6-7) and Song, Zhu
and Chen (1999, figures 222F, 2231), the male paratypes from the same locality with female holo-
type may be mistakenly matched.
DESCRIPTION.— See Zhu and Wang (1991). Chelicerae with three promarginal and two retro-
marginal teeth. Female epigynum without epigynal teeth; epigynal hoods situated posteriorly near
epigastric margin; atrium large, longitudinally elongated; epigynum with anteriorly situated cave.
DISTRIBUTION.— China (Sichuan) (Map 28).
MATERIAL EXAMINED.— None.
Platocoelotes kailiensis Wang, sp. nov.
Figures 77A—E; Map 28
TyprEs.— Holotype male from Kaili, Guizhou, China (X.P. Wang; October 3, 1997), 1 female
paratype from Shanchahe Cave, Maolai Natural Reserve, Libo, Guizhou, China (X.P. Wang;
October 6, 1997), 1 female paratype from WongAng Cave, Maolai Natural Reserve, Libo,
Guizhou, China (X.P. Wang; October 9, 1997), deposited in IZB.
DIAGNOosIs.— The female of this new species is similar to P. icohamatoides but can be distin-
guished by the long, looped copulatory ducts (with at least 5 loops) (Figs. 77A—B). The male can
be distinguished by the posterior extension of the embolic base and the strongly expanded cymbial
furrow (Figs. 77C-E).
FEMALE.— Total length 6.47. Carapace 3.19 long, 2.15 wide. Promargin of chelicera with
three teeth, retromargin with two. Eye sizes and interdistances: AME 0.10, ALE 0.17, PME 0.15,
PLE 0.16; AME-AME 0.08, AME-ALE 0.06, PME-PME 0.09, PME-PLE 0.16, ALE-PLE 0.05,
AME-PME 0.14. Leg measurements: I: 12.3 (3.28, 4.18, 2.91, 1.92); II: 10.4 (2.68, 3.44, 2.60,
1.72); Ill: 9.39 (2.51, 2.89, 2.52, 1.47); IV: 12.6 (3.22, 4.05, 3.50, 1.78). Female epigynum with-
out teeth; epigynal hoods situated posteriorly near epigastric furrow; atrium longitudinally elongat-
ed; epigynum with anterior cavity; copulatory ducts short; spermathecal heads small; spermathecal
bases small, contiguous, longitudinally extending; spermathecal stalks strongly elongated, slender,
highly convoluted with at least 5 loops (Figs. 77A-B).
MALE.— Total length 8.47. Carapace 3.90 long, 2.67 wide. Promargin of chelicera with three
teeth, retromargin with two. Eye sizes and interdistances: AME 0.18, ALE 0.19, PLE 0.19; PME
0.19, AME-AME 0.08, AME-ALE 0.05, PME-PME 0.13, PME-PLE 0.15. Leg measurements: I:
17.5 (4.60, 5.79, 4.42, 2.70); II: 15.5 (4.22, 4.90, 3.91, 2.44); Il: 14.1 (3.75, 4.18, 3.93, 2.19); IV:
18.4 (4.70, 5.61, 5.61, 2.49). Palp with two patellar apophyses; RTA long, occupying almost entire
tibial length; lateral tibial apophysis broad; cymbial furrow elongate, at least 2/3 cymbial length;
conductor broad, with a posteriorly extending apophysis and an anteriorly extending finger-like
apophysis; conductor dorsal apophysis broad; conductor basal lamella small; embolus posterior in
origin; embolic base extending posteriorly; without median apophysis (Figs. 77C-—E).
DISTRIBUTION.— China (Guizhou) (Map 28).
OTHER MATERIAL EXAMINED.— None.
Platocoelotes lichuanensis (Chen and Zhao, 1998)
Figures 78A—B; Map 28
Coelotes lichuanensis Chen and Zhao, 1998:3, figs. 1(1-3) (male holotype from Lichuan, Hubei, China, in
HUW, examined.
Platocoelotes lichuanensis: Wang, 2002:122.
564 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DIAGNOsIS.— The male of this species is similar to P. impletus but can be recognized by the
large lateral tibial apophysis and the relatively strong conductor apical apophyses (Figs. 78A—B).
DESCRIPTION.— Described by Chen and Zhao (1998). Male palp with two patellar apophyses;
RTA long, occuping almost entire tibial length; lateral tibial apophysis broad; cymbial furrow elon-
gate, at least half cymbial length; conductor broad, with a posteriorly extending apophysis and an
anteriorly extending finger-like apophysis; conductor dorsal apophysis broad, lamella small; embo-
lus posterior in origin; embolic base extending prolaterally; without median apophysis (Figs.
78A—B). Female unknown.
DISTRIBUTION.— China (Hubei) (Map 28).
MATERIAL EXAMINED.— CHINA: Hubei: Lichuan, September 21, 1977, male holotype (HUW,
77-598).
Genus Spiricoelotes Wang, 2002
Spiricoelotes Wang, 2002:129 (type species, by original designation, Coelotes zonatus Peng and Wang, 1997
from China).
DIAGNOsIS.— The female can be easily recognized by the absence of epigynal teeth and the
long, strongly convoluted spermathecae, and the male by the strongly curved patellar apophysis,
the elongated cymbial furrow, the absence of a conductor dorsal apophysis, and the slender, ante-
riorly extending, spiraled conductor (Figs. 79-80).
PHYLOGENETIC PLACEMENT.— The absence of both epigynal teeth and median apophysis sup-
ports the sister group relationship between Platocoelotes and Spiricoelotes, together they remain
unresolved with the genera Draconarius, Asiacoelotes, and Bifidocoelotes (Wang, 2002).
DESCRIPTION.— See Wang (2002)
DISTRIBUTION.— China, Japan (Map 29).
COMPOSITION.— 3 species, including | new species:
1. Spiricoelotes urumensis (Shimojana, 1989)
2. Spiricoelotes zonatus (Peng and Wang, 1997)
Coelotes laoyingensis Chen and Zhao, 1997
3. Spiricoelotes pseudozonatus Wang, sp. nov.
Key to the Species of the Genus Spiricoelotes
MMM TALS es See ae. 5 icy e an evSnes cabo vntere aa, sieidvekectys Sallcousy@ioeSs= Lore ee Gey So ene 2
Male (hose'of S. pseudozonatus unknown). .:.....- 3.24. 22.0+ +2. +4 eee 4
2. Spermathecal stalks looped surrounding copulatory ducts (Fig. 80B) .................-- 3
Spermathecal stalks not looped surrounding copulatory ducts (Fig. 79B) ..... pseudozonatus
3. Spermathecal bases extending medially; spermathecal stalks situated close together (Fig. 80B)
Saray eUete ca fUeiisesusils, ifs ay epsieh sl o'e adie ew 'sisei% 2 ceed @Geeyplet ye ocala ne eee rr zonatus
Spermathecal bases extending anteriorly; spermathecal stalks widely separated. . . . urumensis
4. Conductor long, almost reaching distal end of cymbium....................-. urumensis
Conductor short, only reaching half way to distal end of cymbium (Figs. 80C-E) .. . . zonatus
WANG: EAST ASIA COELOTINE SPIDERS 565
Spiricoelotes pseudozonatus Wang, sp. nov.
Figures 79A—B; Map 29
Typrs.— Female holotype from Bao-guang monastery, Chengdu, Sichuan, China (May 21,
1989; P. Beron), deposited in AMNH (Coll. Deeleman).
EtyMOLoGy.— The specific name refers to its similarity to S. zonatus.
DIAGNOsIS.— The female of this new species can be distinguished from S. zonatus by the less
apparent epigynal hoods, the small, medially situated atrium, and the differences in their spermath-
ecal loops (Figs. 79A—B).
FEMALE.— Total length 3.31. Carapace 1.27 long, 0.94 wide. Abdomen 2.04 long, 1.45 wide.
Eye sizes and interdistances: AME 0.05, ALE 0.07, PME 0.06, PLE 0.07; AME-AME 0.02, AME-
ALE 0.02, PME-PME 0.07, PME-PLE 0.07. Leg measurements: I: 3.46 (1.04, 1.25, 0.71, 0.46); II:
3.76 (1.04, 1.30, 0.88, 0.54); Ill: 2.88 (0.82, 0.95, 0.69, 0.42); IV: 3.83 (1.06, 1.31, 0.88, 0.58).
Promargin of chelicera with three teeth, retromargin five to six. Epigynal teeth absent; atrium
small, medially situated; epigynal hoods less apparent; copulatory ducts short; spermathecal heads
not visible; spermathecal bases widely separated; spermathecal stalks strongly elongated, convo-
luted (Figs. 79A—B).
MALe.— Unknown.
DISTRIBUTION.— China (Sichuan) (Map 29).
OTHER MATERIAL EXAMINED.— None.
Spiricoelotes urumensis (Shimojana, 1989)
Map 29
Coelotes urumensis Shimojana, 1989:79, figs. 24-29 (male holotype, male and female paratypes from Ryukyu
Is., in NSMT, not examined).
Spiricoelotes urumensis: Wang, 2002:131.
DIAGNOsIS.— This species is similar to S$. zonatus but can be distinguished by the widely sep-
arated, slender spermathecal tubes of female, and the longer conductor of male.
DESCRIPTION.— See Shimojana (1989). Cheliceral teeth unknown. Female lacking epigynal
teeth; atria small, widely separated; epigynal hoods well developed, situated anterior of atria; cop-
ulatory ducts anteriorly expanded; spermathecal bases widely separated, anteriorly extending; sper-
mathecal stalks long, slender, widely separated, looped around copulatory ducts (Figs. A-B). Male
palp with patellar apophysis long, with distal end sharply curved dorsally; RTA long; lateral tibial
apophysis small; cymbial furrow about half of cymbial length; conductor long, slender, anteriorly
extending; conductor dorsal apophysis absent; conductor lamella small; embolus posterior in ori-
gin, long; median apophysis absent.
DISTRIBUTION.— Ryukyu Is. (Map 29).
MATERIAL EXAMINED.— None.
Spiricoelotes zonatus (Peng and Wang, 1997)
Figures 80A-E; 97]; Map 29
Coelotes zonatus Peng and Wang, 1997:331, figs. 32-36 (2 male and 2 female paratypes, holotype not indi-
cated, from Changsha, Hunan, China, in HBI, examined).— Song, Zhu and Chen, 1999:376, figs. 220H-I,
i jiph, SALE) Wy,
Coelotes laoyingensis Chen and Zhao, 1997:89, figs. 5-6 (female holotype and 1 female paratype from
566 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Laoying, Wudangshan, Hubei, China, in HUW, examined).— Song, Zhu and Chen, 1999:388, figs. 2260,
P, 227P, 229B.
Spiricoelotes zonatus: Wang, 2002:131.
DIAGNOSIS.— This species is similar to S. urumensis but can be distinguished by the broad
spermathecal tubes of female (Figs. AB), and the relatively short conductor of male (Figs. C-E).
DESCRIPTION.— Described by Peng and Wang (1997) and Wang (2002). Chelicerae with five
promarginal and five retromarginal teeth. Female epigynum without epigynal teeth; atria small,
widely separated; epigynal hoods well developed; copulatory ducts short; spermathecal heads
small; spermathecal bases situated close together, medially extending; spermathecal stalks long,
slender, situated close together, looped around copulatory ducts (Figs. A-B). Male palp with patel-
lar apophysis long, with distal end sharply curved dorsally; RTA long; lateral tibial apophysis
small; cymbial furrow longer than half of cymbial length; conductor long, slender, anteriorly
extending; conductor dorsal apophysis absent; conductor lamella small; embolus posterior in ori-
gin, long; median apophysis absent (Figs. C—E).
DISTRIBUTION.— China (Hubei, Hunan, Jiangsu, Sichuan) (Map 29).
MATERIAL EXAMINED.— CHINA: Hunan: Changsha, January 7, 1985, 2 male and 2 female
paratypes (J.F. Wang, HBI); Changsha, Yuelushan, June 1995, 1 female (X.P. Wang, IZB).
Sichuan: Chengdu, Bao-guang Monstery, May 21, 1989, 2 males (P. Beron, Coll. Delleman).
Jiangsu: Nanjing, Zijin (Purple) Mt., 350—450 m, October 9, 1988, 1 female (P. Beron, Coll.
Delleman). Hubei: Wudangshan, Laoying, May 10, 1982, female holotype and | female paratype
of Coelotes laoyingensis (HUW).
Genus Tegecoelotes Ovtchinnikov, 1999
Tegecoelotes Ovtchinnikov, 1999:68 (type species, by original designation, Coelotes bicaudatus Paik, 1976,
from Korea).— Wang, 2002:133.
DIAGNOSIS.— The female can be distinguished by the broad epigynal teeth, the reduced atri-
um, and the short copulatory ducts. The male can be recognized by the elongated patella (except in
T; muscicapus and T: michikoae) and the not spoon-like median apophysis (Figs. 81-84).
Chelicerae with three promarginal and three retromarginal teeth.
PHYLOGENETIC PLACEMENT.— Remains unresolved with Leptocoelotes and the clade with
spoon-like median apophysis and slender epigynal teeth (Wang 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— China, Japan, Korea, Far eastern Russia (Map 30).
COMPOSITION.— 5 species:
1. Tegecoelotes corasides (BOsenberg and Strand, 1906)
Coelotes corasoides Platnick, 1989
2. Tegecoelotes secundus (Paik, 1971)
Tegecoelotes bicaudatus (Paik, 1976)
Coelotes erraticus Nishikawa, 1983
3. Tegecoelotes ignotus (Bésenberg and Strand, 1906)
Agelena ignota Bésenberg and Strand, 1906
4. Tegecoelotes michikoae (Nishikawa, 1977)
5. Tegecoelotes muscicapus (Bésenberg and Strand, 1906)
WANG: EAST ASIA COELOTINE SPIDERS 567
Key to the Species of the Genus Tegecoelotes
Pema Ca (EMD OSELOL FOOLS, UKMOWI)) 35.5 ber « alaiiest,a2e meee cent oak dcauae aoa of Rene, cre wollen? D,
Remalet(choscsOR us CICAPUS UNKTO WIN). 2 ce: SSRIS 215 OL INT Ts Eee ES HUE LA) OT 4
Sepeaella lions. with one patellar apophysis (Figs.:81D-P) . fe... fine). os oes es ee 3)
Patella short, with two patellar apophyses (Figs. 84A—-C)........... muscicapus, michikoae
3. Conductor apex broad, slightly curved anteriorly (Figs. 82C-F)................. corasides
Conductor apex slender, slightly curved posteriorly (Figs. 81C-F) ............... secundus
4. Epigynal teeth close together, separated by less than their width (Fig. 82A).............. 5)
Epigynal teeth separated by approximatedly twice their width ................. michikoae
RES PCRMAtIeCAaLibaSes Withoul-apOphySes sc Gs. seis woe ee a sitter ee oes 6
Spermathecal bases with long, anteriorly extending apophyses (Fig. 83B) .......... ignotus
BeESIpCHaAthecalmcads: slendem(Pips 825) oes heacioke beat attainment aL corasides
Spe hitatmecalaheads, broad \(ETS.. SUB Ye hepasi te Sons 1 anor anet s Aone ta tien te eee ae secundus
Tegecoelotes secundus (Paik, 1971)
Figures 81A—F; Map 30
Tegenaria secunda Paik, 1971:22, figs. 8-14 (male holotype from Korea, deposited in Kyungpook National
University, Taegu, Korea, not examined).— Paik, 1978:360, figs. 162.1—2.
Coelotes bicaudatus Paik, 1976:81, figs. 3, 13-15 (female holotype from Mt. Ode, Korea, deposited in
Kyungpook National University, Taegu, Korea, not examined).— Paik, 1978:334, figs. 147.1-2.
Coelotes erraticus Nishikawa, 1983:125, figs. 1-6 (male and female types from Mt. KomA—gA-take,
Tazawako-cho, Senboku-gun, Akita, Japan, in NSMT, not examined).—Yaginuma, 1971:93, fig.
82.1-2:— Matsuda, 1986:88, figs. 13-16;—Yaginuma, 1986:151, fig. 80.3;— Wang and Zhu, 1991:5,
figs. 13-16:— Song, Zhu and Chen, 1999:375, figs. 217M-N, 219G, N. (First synonymized by Ovtchin-
nikoy, 1999.)
Tegecoelotes bicaudatus Ovtchinnikov, 1999:68, figs. 16-20 (male first described).— Wang, 2002:134, figs.
375-383. (First Synonymized by Marusik and Koponen, 2000.)
Tegecoelotes secunda: Marusik and Koponen, 2000:56.
DIAGNOsIs.— This species is similar to 7: corasides but can be distinguished by the relative-
ly narrow epigynal teeth, the small copulatory ducts, the differences in shapes between the sper-
mathecal tubes of female (Figs. 81A—B) and the slender, posteriorly curved conductor apex, and
the strongly spiraled median apophysis of male (Figs. 81C-—F).
DESCRIPTION.— Described by Paik (1976). Chelicerae with three promarginal and three retro-
marginal teeth. Female epigynal teeth broad, close together; atrium small; copulatory ducts short;
spermathecal heads large; spermathecae with bases small, stalks broad, short, laterally extending
(Figs. 81A—B). Male palp with patella strongly elongated; patellar apophysis present; RTA long:
lateral tibial apophysis large; cymbial furrow short; conductor long, with slender, slightly curved
apex: conductor dorsal apophysis present; conductor lamella small; embolus short, prolateral in
origin; median apophysis not spoon-like, long, with sharp, strongly spiraled apex and membranous
base (Figs. 81C—F).
DISTRIBUTION.— China (Jilin), Korea, Japan, Russia (Far East) (Map 30).
MATERIAL EXAMINED.— RUSSIA: Far East, S-Primorie, “Kedrovaya Pad” Reservation in a liv-
ing house, June 12 — December 24, 1977, 2 males and 2 females (B.P. Zakharov, SZM). KOREA:
Moon-Kyang-Sae-Jae, 1 male, August 20, 1990 (J.P. Kim, KAI).
568 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Tegecoelotes corasides (Bésenberg and Strand, 1906)
Figures 82A—F; Map 30
Tegenaria corasides Boésenberg and Strand, 1906:301, fig. 459 (fig. 460 is Paracoelotes luctuosus) (male and
female types, holotype not indicated, from Saga, Japan, in SMF, examined).— Yaginuma, 1957:17, fig. 2;
— Yaginuma, 1960:92, fig. 81.8:— Yaginuma, 1971:92, fig. 81.8.
Coras luctuosus: Saito, 1934:342, figs. 27, 53.— Saito, 1959:43, fig. 16A—C (misidentification).
Coelotes modestus: Nishikawa, 1974:177, figs. 17—19.— Nishikawa, 1977: figs. 19-20 (misidentification).
Coelotes corasides: Yaginuma, 1986:148, fig. 80.1.— Chikuni, 1989:103, fig. 21.
Coelotes corasoides: Platnick, 1989:422 (lapsus).
Tegecoelotes corasides: Wang, 2002:134.
DIAGNOSIS.— This species is similar to TZ. secundus but can be distinguished by the relatively
broad epigynal teeth, the large copulatory ducts, the differences in shapes between the spermathe-
cal tubes of female (Figs. 82A—B) and the broad, anteriorly curved conductor apex, and the simple
median apophysis of male (Figs. 82C—F).
DESCRIPTION.— Described by Bésenberg and Strand (1906). Chelicerae with three promargin-
al and three retromarginal teeth. Epigynal teeth broad, close together; atrium small; copulatory
ducts large; spermathecal heads small; spermathecae with bases small, stalks broad, short (Figs.
82A-—B). Male palp with patella strongly elongated; patellar apophysis present; RTA long; lateral
tibial apophysis large; cymbial furrow short; conductor long, with broad, anteriorly curved apex;
conductor dorsal apophysis present; conductor lamella small; embolus short, prolateral in origin;
median apophysis not spoon-like, long (Figs. 82C—F).
DISTRIBUTION.— Japan (Map 30).
MATERIAL EXAMINED.— JAPAN: Saga, 1 male and | female types (W. Donitz, SMF, 4808);
Saga, 2 male and 9 female paratypes (W. Donitz, SMF, 4809); Cross Kamdeals, 1 male and 1
female (Donitz, ZMB, 31192); Kauagawa, February 18, 1905, 1 male (H. Sauter, ZMB,31191);
Camp Fuji, March 3, 1955, 1 male (V. Cambl, CAS).
Tegecoelotes ignotus (Bosenberg and Strand, 1906)
Figure 83A—B; Map 30
Agelena ignota Bésenberg and Strand, 1906:299, fig. 466 (2 female types, holotype not indicated, from Japan,
in SMF, examined).— Ishinoda, 1957:12, fig. 3.
Coelotes ignotus: Lehtinen, 1967:224;— Nishikawa, 1974:178, fig. 33.
Tegecoelotes ignotus: Wang, 2002:134.
DIAGNOsIS.— The female of this species can be easily recognized by the broad, anteriorly sit-
uated epigynal teeth, the broad copulatory ducts, and the long, slender spermathecal diverticula
(Figs. 83A—B).
DESCRIPTION.— Described by Bésenberg and Strand (1906). Chelicerae with three promargin-
al and three retromarginal teeth. Female epigynum with epigynal teeth broad, more or less round-
ed, situated anteriorly; atrium small; copulatory ducts broad, extending laterally; spermathecal
heads not visible; spermathecal bases with long, anteriorly extending diverticula; spermathecal
stalks slender, anteriorly extending (Figs. 83A—B). Male unknown.
DISTRIBUTION.— Japan (Map 30).
MATERIAL EXAMINED.— JAPAN: no detailed label, 2 female types (W. Donitz, SMF, 4697).
WANG: EAST ASIA COELOTINE SPIDERS 569
Tegecoelotes michikoae (Nishikawa, 1977)
Map 30
Coelotes michikoae Nishikawa, 1977:39, figs. 13-18 (female holotype, male and female paratypes from
Minoo, Osaka Prefecture, Japan, deposited in the Osaka Museum of Natural History, Osaka, and in the
Arachnological Society of Japan, Ohtemon-Gakuin University, Osaka, Japan, not examined).— Chikuni,
1977:56, fig. 1.3:— Yaginuma, 1986:151, fig. 80.2;— Chikuni, 1989:102, fig. 20.
Tegecoelotes michikoae: Wang, 2002:134.
DIAGNOsIS.— The male of this species is similar to 7? muscicapus and can only be distin-
guished by the less expanded conductor apex. The widely separated epigynal teeth can distinguish
females from other Tegecoelotes (except T: muscicapus, which is not known).
DESCRIPTION.— See Nishikawa (1977). Chelicerae with three promarginal and three retromar-
ginal teeth. Female epigynal teeth broad, widely separated by at least twice their width; atrium
small; spermathecal bases extending medially; spermathecal stalks situated close together. Male
palp with patella short, approximately tibial length; patella with two apophyses, with dorsal one
small; RTA long; lateral tibial apophysis present; cymbial furrow short; conductor long, with ante-
riorly curved apex; conductor dorsal apophysis present; conductor lamella small; embolus short,
prolateral in origin; median apophysis not spoon-like, with slender apex.
DISTRIBUTION.— Japan (Map 30).
MATERIAL EXAMINED.— None.
Tegecoelotes muscicapus (Bosenberg and Strand, 1906)
Figure 84A—C; Map 30
Tegenaria muscicapa Bosenberg and Strand, 1906:302, fig. 479 (male holotype from Saga, Japan, in SMF,
examined).
Tegecoelotes muscicapa: Wang, 2002:134.
DIAGNOsIs.— The male of this species is similar to 7. michikoae and can only be distinguished
by the relatively broad conductor apex (Figs. 84A—B).
DESCRIPTION.— Described by Bosenberg and Strand (1906). Chelicerae with three promargin-
al and three retromarginal teeth. Male palp with patella short, approximately tibial length; patella
with two apophyses, with dorsal one small; RTA long; lateral tibial apophysis present; cymbial fur-
row short; conductor long, with broad, anteriorly curved apex; conductor dorsal apophysis present;
conductor lamella small; embolus short, prolateral in origin; median apophysis not spoon-like, with
slender apex (Figs. 84A—C). Female unknown.
DISTRIBUTION.— Japan (Map 30).
MATERIAL EXAMINED.— JAPAN: Saga, male holotype (W. Donitz, SMF, 4820); Saga, | male
(W. Donitz, SMF, in the same vial as Coelotes corasides paratypes, 4809).
Genus Tonsilla Wang and Yin, 1992
Tonsilla Wang and Yin, 1992:263 (type species, by original designation, Tonsilla truculenta Wang and Yin,
1992 from China).— Platnick, 1997:671;— Wang, 2002:136.
DIAGNOsIS.— The female can be easily recognized by the large epigynal atrium, the posterior
extension of anterior atrial margin, the median, closely situated epigynal teeth, and the large cop-
ulatory ducts, and the male by the long patellar apophysis, the short cymbial furrow, and the bifur-
570 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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cate (or lobed) conductor (Figs. 85-95). Cheliceral promargin with three teeth, retromargin with
two.
PHYLOGENETIC PLACEMENT.— The presence of a large atrium and the posteriorly extending
anterior atrial margin support the sister group relationship between TJonsilla and Paracoelotes.
Together they are the sister group of the clade with two retromarginal cheliceral teeth and large
conductor lamella (Wang 2002).
DESCRIPTION.— See Wang (2002).
DISTRIBUTION.— Central China (Maps 31-32)
COMPOSITION.— 7 species, including 1 new species and 2 new combinations:
The truculenta group species
1. Tonsilla eburniformis Wang and Yin, 1992
2. Tonsilla imitata Wang and Yin, 1992
3. Tonsilla truculenta Wang and Yin, 1992
The variegates group species
4. Tonsilla lyratus (Wang et al., 1990), NEW COMBINATION
5. Tonsilla tautispinus (Wang et al., 1990), NEw COMBINATION
6. Tonsilla variegatus (Wang et al., 1990)
7. Tonsilla makros Wang, sp. nov.
Key to Species of the Genus Tonsilla
Viale pity Pets pect: bere SR to ee AR UR 8 ee D)
Beri ale a0 Hei ehs eee cdg steine tel betas RL: DEE AIR RES 4
2-Conductor biturcate (Figs. 94C=E). 2 2 oo es) se sce ees eke hs shee et 0 oe oe eee 3
Conductor positenorly: lobedi(Figs::86C-E) (2 -92-- eae oe = eerie ae truculenta
3. Conductor strongly bifurcate, with anterior apophysis slightly curved anteriorly (Figs. 94D, E)
BR eR Co RENCE RP ae ee PRES eae OR SEL ET OCR RE I Ns oo Coe varlegatus
Conductor slightly bifurcate, apex not curved (Figs. 95A-C)................-00-- makros
Epigynal teeth long, slender, close together or separated by less than their width, spermathecae
longitudinally elongate (Figs. 86A; 91A)............... truculenta, eburniformis, imitata
5» Epigynal teeth situated on anterior atrial margin (Fig. 94A):.. 2. .... i. 2a. 2 eee 6
Epigynal teeth situated anterad of atrium, separated from anterior atrial margin (Fig. 93A)
Dea ifort Sissel Yotiatas 8 cal oa yn). nin lw cx ie hie unr tncke Sere an tg oe CRA es cena a ae tautispinus
6. Copulatory ducts strong expanded anteriorly; spermathecal heads situated anteriorly (Fig. 94B)
fet ele She sh Soe Le ee Se EE ee eee variegatus
Copulatory ducts not anteriorly expanded; spermathecal heads situated laterally (Fig. 92B)
Moree SR Os hiss . AE iw oa aa ae ee lyratus
The “truculenta” Group
The male of this group generally has the long, dorsally curved patellar apophysis and posteri-
orly lobed conductor; the female has the slender, closely situated epigynal teeth, and anteriorly
elongated spermathecae.
WANG: EAST ASIA COELOTINE SPIDERS 571
Tonsilla eburniformis Wang and Yin, 1992
Figures 85A—B; Map 31
Tonsilla eburniformis Wang and Yin, 1992:265, figs. 13-14 (female holotype from Muyu, Shenlongjia, Hubei,
China, in HTU, examined).
DraGnosis.— The female of this species is similar to T. truculenta and can be distinguished
by the slightly separated bases of epigynal teeth, the broad septum, and the short, broad spermath-
ecae (Figs. 85A—B).
DESCRIPTION.— Described by Wang and Yin (1992). Cheliceral promargin with three teeth,
retromargin with two. Female epigynal teeth slender, situated close together, with bases slightly
separated; atrium large; septum broad; copulatory ducts situated mesad of spermathecae, slightly
extending anteriorly; spermathecal heads long, extending mesad of spermathecae; spermathecae
broad, widely separated (Figs. 85A—B). Male unknown.
DISTRIBUTION.— China (Hubei) (Map 31).
MATERIAL EXAMINED.— CHINA: Hubei: Shenlongjia, Muyu, November 12, 1992, female
holotype (J.F. Wang, HTU).
Tonsilla imitata Wang and Yin, 1992
Map 31
Tonsilla imitata Wang and Yin, 1992:264, figs. 11-12 (female holotype from Qiayang, Hunan, China, in HBI,
not examined).— Song, Zhu and Chen, 1999:395, figs. 230G—H.
DiAGNosis.— The female of this species is similar to T. truculenta (specimen from Mitai,
Guizhou, variation 5) and can only be distinguished by the slightly different epigynal teeth.
DESCRIPTION.— See Wang and Yin (1992). Cheliceral promargin with three teeth, retromar-
gin with two. Female epigynal teeth slender, situated close together; atrium large; copulatory ducts
situated mesad of spermathecae, slightly extending anteriorly; spermathecal heads long, originat-
ing mesad of spermathecae, situated anteriorly; spermathecae broad, widely separated. Male
unknown.
DISTRIBUTION.— China (Hunan) (Map 31).
MATERIAL EXAMINED.— None.
Tonsilla truculenta Wang and Yin, 1992
Figures 86A—E; Map 31
Tonsilla truculenta Wang and Yin, 1992:263, figs. 1-10 (female holotype, 2 male and 7 female paratypes from
Tianzishan, Sangzhi, Hunan, China, in HTU, examined).— Song, Zhu and Chen, 1999:395, figs. 131,
230I-L:— Wang, 2002:137, figs. 384-399.
DIAGNOsIs.— The female can be recognized by the slender, closely situated epigynal teeth, the
broad atrium, the broad copulatory ducts, and the long spermathecae heads (Figs. 86A—B). The
male can be distinguished by the long, strongly curved patellar apophysis and the presence of a
lobed conductor (Figs. 86C—E).
DESCRIPTION.— Described by Wang and Yin (1992). Cheliceral promargin with three teeth,
retromargin with two. Female epigynal teeth slender, situated close together, more or less over-
lapped; septum broad; copulatory ducts broad, situated mesad of spermathecae, not anteriorly
extending; spermathecal heads long; spermathecae longitudinally extending, widely separated
SP PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
(Figs. 86A—B). Male palp with patellar apophysis long, dorsally curved; RTA long; lateral tibial
apophysis present, widely separated from RTA; cymbial furrow short; conductor lobed; conductor
dorsal apophysis slender, with apex slightly toothed; conductor lamella small; median apophysis
spoon-like, elongated (Figs. 86C-E).
DISTRIBUTION.— China (Guizhou, Hunan, Sichuan) (Map 31).
MATERIAL EXAMINED.— CHINA: Hunan: Sangzhi, Tianzishan, October 27, 1989, female holo-
type, male allotype, | male and 7 female paratypes (J.F. Wang, HTU).
VARIATIONS.— Further collection and examination of this species shows strong variation in
both male and female genitalia, which includes the size and shape of patellar apophysis, the shape
of conductor, and the shape of conductor dorsal apophysis of male; and the shape and position of
epigynal teeth, the size and shape of septum, the atrial shape, the size and shape of spermathecae,
and the shape of copulatory ducts of female. It is likely that the species 7: eburniformis and T. imi-
tata also fall into those variations. Further collections of male specimens from above two species
localities are needed.
VARIATION |. CHINA: Hunan: Sangzhi, Tianzishan, October 27, 1989, 6 male and 2 female
paratypes of 7: truculenta (J.F. Wang, HTU). From the same locality with female holotype and male
allotype, but differs by the elongated patellar apophysis, the small conductor lobe, the sharp, none
toothed apex of conductor dorsal apophysis of male (Figs. 87C—D) and by the slender septum and
the anteriorly extending, bifurcate copulatory ducts of female (Figs. 87A—B).
VARIATION 2. CHINA: Hunan: Sangzhi, Tianzishan, October 27, 1989, 1 female paratype of T.
truculenta (J.F. Wang, HTU). From the same locality with female holotype and male allotype, but
differs by the different atrial shape, the anterially extending, broad copulatory ducts, and the short,
broad spermathecae of female (Figs. 88A—B).
VARIATION 3. CHINA: Sichuan: Chunging, Jiyunshan, October 26, 1997, 2 females and 1 male
(X.P. Wang, HTU). Differs by the short patellar apophysis, the broad conductor lobe, and the slight-
ly bifurcate conductor dorsal apophysis of male (Figs. 89C—E) and the relatively strong epigynal
teeth, the slightly anterior extension of copulatory ducts, the short, broad spermathecae of female
(Figs. 89A-B).
VARIATION 4. CHINA: Guizhou: Guiyang, campus of Guizhou Teachers University, October 30,
1997, 1 male (X.P. Wang, HTU). Differs by the blunt conductor apex, the large conductor lobe, and
the short, sharply pointed patellar apophysis of male (Figs. 90A—B).
VARIATION 5. CHINA: Guizhou: Meitan, August 1981, 1 female paratype of 7: truculenta (F.J.
Li, HTU). Hunan: Chengbu, August 21, 1982, | female paratype of T. truculenta (J.F. Wang,
HTU). Differs by the unique copulatory ducts, the short, broad spermathecae (Fig. 91A—B, female
paratype from Meitan, Guizhou).
The “variegatus” Group
The male of this group typically has the strongly elongated patellar apophysis (as long as patel-
lar length) and non-lobed, more or less bifurcate conductor; the female has the short, slightly sep-
arated epigynal teeth and short spermathecae.
Tonsilla lyratus (Wang et al., 1990), NEW COMBINATION
Figures 92A—B; 97H; Map 32
Coelotes lyratus Wang et al., 1990:200, figs. 55-56 (female holotype from Tianpingshan, Sangzhi, Hunan
China, in HBI, examined).— Song, Zhu and Chen, 1999:376, figs. 220T—U.
DIAGNOsISs.— The female of this species is similar to 7. tautispinus but can be distinguished
WANG: EAST ASIA COELOTINE SPIDERS 513
by the closely situated epigynal teeth and atrium, the anteriorly situated spermathecal heads, and
the anterior extension of spermathecae (Fig. 92A—B).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth situated close together, slightly separated from anterior
atrial margin; atrium large; epigynal hoods posteriorly situated, near epigastric furrow; copulatory
ducts broad, situated mesad of spermathecae; spermathecal heads anteriorly situated; spermathecae
broad, anteriorly extending, slightly zig-zaged (Figs. 92A—B). Male unknown.
DISTRIBUTION.— China (Hunan) (Map 32).
MATERIAL EXAMINED.— CHINA: Hunan: Sangzhi, Tianpingshan, October 16, 1986, female
holotype (J.F. Wang, HBI).
Tonsilla tautispinus (Wang et al., 1990), NEW COMBINATION
Figures 93A—B; Map 32
Coelotes tautispinus Wang et al., 1990:190, figs. 32-33 (female holotype from Lushan, Jiangxi, China, in
HBI, examined).— Song, Zhu and Chen, 1999:378, figs. 225K—L.
DIAGNOsIS.— The female of this species is similar to 7. lyratus but can be distinguished by the
widely separated epigynal teeth and atrium, the laterally situated spermathecal heads, and the lat-
eral extension of spermathecae (Fig. 93A—B).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth close together, anteriorly situated, wide apart from atrium;
atrium large; epigynal hoods medially situated; copulatory ducts broad, situated anterad of sper-
mathecae; spermathecal heads laterally originating and extending; spermathecae broad, short,
slightly extending laterally (Figs. 93A—B). Male unknown.
DISTRIBUTION.— China (Jiangxi) (Map 32).
MATERIAL EXAMINED.— CHINA: Jiangxi: Lushan, June 15, 1987, female holotype (J.F. Wang,
HBI).
Tonsilla variegatus (Wang et al., 1990)
Figures 94A—F; Map 32
Coelotes variegatus Wang et al., 1990:184, figs. 20-24 (female holotype and | male paratype from Huang-
shan, Anhui, China, in HBI, examined).— Song, Zhu and Chen, 1999:388, figs. 226E—-F, 227N, 2280.
Tonsilla variegatus: Wang, 2002:136.
DIAGNOsIs.— The male of this species is similar to 7: makros but can be distinguished by the
slightly spiraled, strongly bifurcate conductor and the relatively long median apophysis (Figs.
94C_-F). The female can be recognized by the broad atrium, the large copulatory ducts, the poste-
riorly originated spermathecal heads, and the small spermathecae (Figs. 94A—B).
DESCRIPTION.— Described by Wang et al. (1990). Cheliceral promargin with three teeth, retro-
margin with two. Female epigynal teeth short, close together, situated anteriorly close to anterior
atrial margin; atrium large; copulatory ducts broad, strongly expanded anteriorly; spermathecal
heads long, originating posteriorly and extending laterally; spermathecae small (Figs. 94A—B).
Male palp with patellar apophysis strongly elongated, at least the patellar length; RTA long; later-
al tibial apophysis large; cymbial furrow short; conductor strongly bifurcate, with slightly spiraled
apex; conductor dorsal apophysis slender, lamella small; embolus prolateral in origin; median
apophysis spoon-like, transversely elongated (Figs. 94C—F).
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Volume 54, No. 26
DISTRIBUTION.— China (Anhui) (Map 32).
MATERIAL EXAMINED.— CHINA: Anhui: Huangshan, October 27, 1974, female holotype and
male paratype (J.F. Wang and C.M. Yin, HBI).
Tonsilla makros Wang, sp. nov.
Figures 95A—C; Map 32
Types.— Male holotype and male paratype from Wong-Ang, Libo Co., Guizhou, China
(October 9, 1997; X.P. Wang and J.C. Ran), deposited in AMNH (holotype), IZB (1 male paratype).
ETYMOLOGY.— The specific name refers to the long patellar apophysis.
DIAGNOSIS.— The male of this species is similar to 7. variegatus but can be distinguished by
the slightly bifurcate conductor, and the relatively short median apophysis (Figs. 95A—C).
MALES.— Total length 6.20. Carapace 2.40 long, 2.80 wide. Cheliceral promargin with three
teeth, retromargin with two. Eye sizes and interdistances: AME 0.08, ALE 0.15, PME 0.14, PLE
0.15, AME-AME 0.05, AME-ALE 0.02, PME-PME 0.04, PME-PLE 0.06, ALE-PLE 0.02, AME-
PME 0.08. Leg measurements: I: 7.20 (1.84, 2.48, 1.80, 1.08); II: 6.20 (1.80, 2.08, 1.52, 0.80); III:
5.60 (1.68, 1.72, 1.48, 0.72); IV: 8.00 (2.08, 2.48, 2.44, 1.00). Male palp with patellar apophysis
strongly elongated; RTA long; lateral tibial apophysis large, widely separated from RTA; cymbial
furrow short; conductor slightly bifurcate; conductor dorsal apophysis slender; conductor lamella
small; embolus prolateral in origin; median apophysis spoon-like, short (Figs. 9sA—C).
FEMALE.— Unknown.
DISTRIBUTION.— China (Guizhou) (Map 32).
OTHER MATERIAL EXAMINED.— None.
COELOTINAE DISTRIBUTION PATTERNS
Species of the spider subfamily Coelotinae are collected from North America (west to the
Rocky Mountains), Europe, Central Asia, the Himalayas, and East Asia (south to Nepal and north-
ern Vietnam). The single species recorded from Lebanon (Coelotes coedatus de Blauwe, 1973)
could be a mistake because no further report of coelotines from this area so far. Coelotines are high-
ly endemic species. No genus is shared between North America and Eurasia. Two genera (Coelotes
and Paracoelotes) are found across Eurasia, but no species is shared between Europe and East Asia.
Most species are endemic to small-restricted ranges, especially those from the Himalayas and East
Asia, and only few are quite widespread and have large, but limited ranges (not beyond the bound-
ary of Europe, Central Asia, the Himalayas, or East Asia), judging from the examined collections
and available publications. East Asia is not only the richest in genera by having at least 15 (with
12 unique genera), it’s also the richest in species (Table 1). Of the twenty coelotinae genera, only
three of them are widely distributed across Eurasian continent (Table 2). Central Asia holds species
from all three widespread genera, but lacks its own unique genus.
The genera Coras and Wadotes, with 15 and 11 species separately, are endemic to North
America. They are collected from southeast Canada (southern Ontario, Southern Quebec, New
Brunswick, Nova Scotia, and Newfoundland) and eastern US (west to Minnesota, Iowa, Kansas,
Oklahoma, and Texas; south to northern Florida). The occurrence of one male Coras lamellosus
(Keyserling, 1887) from Medicine Hat, Alberta suggests that C. lamellosus might extend its distri-
bution west to southern Manitoba, southern Saskatchewan, and possibly North Dakota too, but this
collection need to be verified.
The genera Eurocoelotes and Urocoras, with 11 and five species separately, are recorded from
WANG: EAST ASIA COELOTINE SPIDERS SS)
TABLE |. Numbers of Coelotinae genera and species distributed in North America, Europe, Central Asia,
the Himalayas, and East Asia
North America Europe Central Asia The Himalayas East Asia
# of genera 2 4 3 2 15
Unique genera 2 2 0 1 12
#f of species? 26 48 19 14 (>30 undescribed) >200 (including some
undescribed)
2 All data from Platnick (2000-2002) and Wang (2002)
3 The published data, maybe slightly different
TABLE 2. Three widespread Coelotinae genera*
North America — Europe Central Asia The Himalayas East Asia
Coelotes = yes yes — yes
Draconarius — — yes yes yes
Paracoelotes _ yes yes - yes
4 All data from Platnick (2000-2002) and Wang (2002)
Europe. Most species are known from eastern and southeastern Europe (Italy, Greece, Hungary,
Bulgaria, former Yugoslavia, and Turkey), and only E. inermis (L. Koch, 1855) is widespread from
France, Germany, Poland, Switzerland, Italy, Austria, former Yugoslavia, to Bulgaria. Turkey has
the richest Urocoras species diversity (currently two species, U. nicomedis (Brignoli, 1978) and U.
phthisicus (Brignoli, 1978), and four more from author’s unpublished data).
Sister to Urocoras is the Himalcoelotes, which is exclusively limited to the region of the
Himalayas. All 10 species are recorded from Nepal, including at least 2 species that extended their
distribution to the Tibet side of the Himalayas.
Two genera, Coelotes and Paracoelotes, are widely distributed across Eurasia. Coelotes, here
refers to only those species that belong to the type species clade (C. atropos Walckenaer, 1830),
includes groups (atropos, charitonovi, exitialis, and pseudoterrestris) defined by Wang (2000).
The atropos group species are found from Europe, Central Asia to Xinjiang, western China, and
the charitonovi group species are only limited to Central Asia. The pseudoterrestric group species
from Yunnan, China and the exitialis group species from Japan have quite allopatric distribution
patterns compared with atropos and charitonovi groups. Paracoelotes, with 17 species, is found in
Europe, Central Asia, and East Asia. Two Paracoelotes species are widespread in East Asia.
According to the specimens examined in this study, P. spinivulvus is one of the common species in
northern China (Jilin, Beijing, Hebei, Shanxi, Shaanxi, Gansu, and Hubei), Japan (no locality
label), Korea, and Far East Russia, and another common species, P. luctuosus, is distributed in
southern China (Hubei, Guizhou, Zhejiang, Jiangsu, Anhui, and Sichuan) and Japan (Osaka, Kobe,
Yokohama, Tokyo, and Saga). The species P. spinivulvus and P. luctuosus are very similar in gen-
italic morphology and they distributional regions overlap in central China. Both of them were col-
lected from Hubei (Xiangfan) and Guizhou (Guiyang) by the author.
Sister to Paracoelotes is the Tonsilla, which has seven species and is widespread in central and
eastern China.
Both Ambanus and Tegecoelotes have limited distributions in northeast regions of East Asia.
Ambanus, with 18 species, occurs in Korea, Far East Russia, and northeast China, while
576 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. X
Tegecoelotes, with five species, is mostly found in Japan. Only one Tegecoelotes species is found
widespread in Japan, northeast China, Korea, and Far East Russia.
Sister to Ambanus is Robusticoelotes, which has only one species, from eastern China
(Zhejiang and Jiangsu).
The genus Femoracoelotes, with two species and uniquely identified by the presence of a
femoral apophysis, is found only in Taiwan Island. Together with the sister genus Coronilla (with
five species) from central and eastern China and northern Vietnam they form the sister group of all
other coelotine clades.
Asiacoelotes, with 15 species, occurs throughout East Asia (Japan, Korea, and China, west to
Hunan and Guangdong provinces).
Spiricoelotes, with only three species, occurs from central and eastern China to Ryukyu Island
and is sister to Platocoelotes, which has five central Chinese species.
Draconarius, with 86 species, is the most specious genus and is widespread from the
Himalayas to central and eastern China. The type species, D. venustus from Tadzhikistan, is the
only Draconarius species from that region. The closest published distribution sites to D. venustus
are Nepal, Tibet and Gansu (China). Judging from this distribution pattern and the large number of
undescribed species from the Himalayas (by examining Dr. J. Martens collections from his
Himalaya Expeditions), Draconarius occurrence along the Karakoram mountain range and more
Draconarius species in Tadzhikistan region are possible.
The remaining three genera are only recorded from China. Bifidocoelotes, with two species,
from Hong Kong and Taiwan, Leptocoelotes, with two species, from Zhejiang and Taiwan, and
Longicoelotes, with only three species, is recorded from eastern China (Zhejiang, Jiangsu, and
Fuji) and Senkaku.
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Muséum national d’ Histoire naturelle (A, Zool.) 25:1481.
SHIMOIJANA, M. 1989. Four new species of the genus Coelotes (Araneae:Agelenidae) from the Ryukyu Islands,
Japan. Pages 75-82 in Y. Nishikawa, and H. Ono, eds., Arachnological Papers Presented to Takeo
Yaginuma on the Occasion of his Retirement. Osaka Arachnologists’ Group, Osaka, Japan.
SHIMOJANA, M. 2000. Description of eleven new species of the genus Coelotes (Araneae: Amaurobiidae) from
the Ryukyu Islands, Japan. Acta Arachnologica Tokyo 49:165—189.
SonG, D.X., M.S. ZHU AND J. CHEN. 1999. The Spiders of China. Hebei Science and Technology Publishing
House, Shijiazhuang, China. 640 pp.
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WANG, J.F., C.M. YIN, X.J. PENG, AND L.P. Xt. 1990. New species of the spiders of the genus Coelotes from
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WANG, J.F. 1994. Descriptions of a new genus and two new species of Amaurobiidae from China (Araneae).
Acta Zootaxonomica Sinica 19:281—285.
WANG, J.F., AND C.D. ZHu. 1991. Four new species and a new record of the genus Coelotes from China
(Araneae:Agelenidae). Sichuan Journal of Zoology 10(4):3-6.
WANG, J.F., AND C.M. YIN. 1992. A new genus and three new species of funnel-web spiders from south China
(Araneae: Agelenidae). Journal of the Hunan Normal University (Natural Science) 15:263—272.
WANG, L. AND Y.J. Xu. 1988. A new species of Coelotes from China (Araneae, Agelenidae). Journal of
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WANG, X.P., AND H. ONo. 1998. The coelotine spiders (Araneae,Amaurobiidae) of Taiwan. Bulletin of the
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WANG, X.P., I. Tso, AND H.Y. Wu. 2001. Three new Coelores spiders (Araneae, Amaurobiidae) from Taiwan.
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WANG, X.P. 2002. A generic-level revision of the spider subfamily Coelotinae (Araneae, Amaurobiidae).
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YAGINUMA, T. 1960. Spiders of Japan in Color. Hoikusha Publishing Co., Osaka. 186 pp.
YAGINUMA, T. 1971. Spiders of Japan in Color (enlarged and revised ed.). Hoikusha Publishing Co., Osaka.
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YAGINUMA, T. 1986. Spiders of Japan in Color (new ed.).Hoikusha Publishing Co., Osaka, Japan. 350 pp.
ZHANG, Y.J., X J. PENG, AND J.P. Kim. 1997. Five new species of the genus Coelotes (Araneae:Agelenidae)
from China. The Korean Journal of Systematic Zoology 13(4):291—296
ZHANG, Y J. AND C.M. Yin. 2001. A new species of the genus Coelotes from China (Araneae:Amaurobiidae).
Acta Zoologica Sinica 10:11—12.
ZHANG, Z.S., M.S. ZHU AND D.X. SONG. 2002. Three new species of the subfamily Coelotinae from Mt.
Shennongjia of Hubei province, China (Araneae:Amaurobiidae). Journal of Baoding Teacher's College
WANG: EAST ASIA COELOTINE SPIDERS SS)
15:52-55.
Zuu, C.D. AND J.F. WANG. 1991. Six new species of the genus Coelotes from China (Araneae:Agelenidae).
Journal of Norman Bethune University Medical Sciences 17(5):1+4.
Zuu, C.D. AND J.F. WANG. 1994. Seven new species of the genus Coelotes from China (Araneae:Agelenidae).
Acta Zoologica Sinica 19:37-45.
580
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ILLUSTRATIONS
AND
DISTRIBUTION MAPS
WANG: EAST ASIA COELOTINE SPIDERS 58}
spermathecalhead ———
Wil ee.
“) = epigynal tooth SY
AOD (eee
Ae Ph (4).
. , ; 5 iy ; 3 : ‘ ‘ f* 7
Ax) spemmathecas —- (7
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= VW Ae
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AN S Sd Ga 2 Se OTE Halles : e i: | 7
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Ficure 1. Bifidocoelotes bifidus (Wang, Tso and Wu). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D.
Pedipalpus, retrolateral view.
FiGURE 2. Bifidocoelotes primus (Fox), female. A. Epigynum. B. Vulva.
582 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 3. Coronilla gemata Wang. A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus, ventral view;
E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 583
conductor dorsal apophysis
embolus
<\\) conductor 7e Sa
US
UE B
patellar apophyses
cymbial furrow
FicurE 4. Coronilla libo Wang, sp. nov. A. Pedipalpus, prolateral view, B. Pedipalpus, ventral view. C. Pedipalpus,
retrolateral view.
584 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Ficure 5. Coronilla sigillata Wang. A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus, ventral view.
E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 585
FiGuRE 6. Coronilla subsigillata Wang, sp. nov. A. Epigynum. B. Vulva.
epigynal hood atrium sperrmathecal head
copulatory duct
conductor dorsal apophysis
conductor (trifid)
cymbial furrow
lateral tibial apophysis—2
patellar apophysis \ “og
FiGuRE 7. Draconarius absentis Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
586 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGureE 9. Draconarius adligansus (Peng and Yin). A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 587
Ficure 10. Draconarius agrestis Wang, sp. nov. A. Pedipalpus, ventral view. B. Pedipalpus, retrolateral view.
588 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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/
Se
ackien
i
/
Hl
FicurE 12. Draconarius argenteus Wang, sp. nov. A. Pedipalpus, ventral view. B. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 589
FIGURE 13. Draconarius aspinatus (Wang et al.). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus,
ventral view. E. Pedipalpus, retrolateral view.
590 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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4 \ : Be
A 5
FiGurE 14. Draconarius baronii (Brignoli). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view. C. Pedipalpus,
retrolateral view.
FiGureE 15. Draconarius baxiantaiensis Wang, sp. nov. A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS ay)
Figure 16. Draconarius bituberculatus (Wang et al.). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view. C.
Pedipalpus, retrolateral view.
Aas
Figure 17. Draconarius calcariformis (Wang). A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
592 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 19. Draconarius carinatus (Wang et al.). A. Epigynum. B. Vulva. C. padipalpus, prolateral view. D. Pedipalpus,
ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 393
FIGURE 20. Draconarius chaigiaoensis (Zhang, Peng and Kim). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral
view. C. Pedipalpus, retrolateral view.
FIGURE 21. Draconarius cheni (Platnick). A. Vulva. B. Epigynum.
594 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 22. Coelotes coreanus (Paik and Yaginuma). A. Epigynum. B. Vulva. C. pedipalpus, prolateral view. D. Pedi-
palpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 55
FiGuRE 23. Draconarius curiosus Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. C. Pedipalpus,
retrolateral view.
596 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 24. Draconarius davidi (Schenkel). A. Epigynum. B. Vulva.
FIGURE 25. Draconarius denisi (Schenkel). A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS
S7/
spe
A
re!
FIGURE 26. Draconarius digitusiformis (Wang et al.). A. Epigynum. B. Vulva. C. padipalpus, prolateral view.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
598 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 27. Draconarius disgregus Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
FIGURE 28. Draconarius dissitus Wang, sp. nov. A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 599
FiGURE 29. Draconarius dubius Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
FiGureE 30. Draconarius episomos Wang, sp. nov. A. Epigynum. B. Vulva.
600 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 31. Draconarius griswoldi Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
FIGURE 32. Draconarius gurkha (Brignoli). A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 601
FIGURE 34. Draconarius hangzhouensis (Chen). A. Epigynum. B. Vulva.
602 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 35. Draconarius incertus Wang, sp. nov. A. Pedipalpus, ventral view. B. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS
FIGURE 37. Draconarius labiatus (Wang and Ono). A. Epigynum. B. Vulva.
603
604 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FicureE 38. Draconarius linxiaensis Wang, sp. nov. A. Pedipalpus, ventral view. B. Pedipalpus, retrolateral view.
605
WANG: EAST ASIA COELOTINE SPIDERS
rius lutulentus (Wang et al.), types. A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D.
FIGURE 39. Dracona
Pedipalpus,
trolateral view.
ventral view. E. Pedipalpus, re
606 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 40. Draconarius lutulentus (Wang et al.), showing genitalic variations:A—D from Wudangshan, Hubei, China;
E-H from Hongping, Hubei, China. A, E. Epigynum. B, F. Vulva. C, G. Pedipalpus, ventral view. D, H. Pedipalpus, retro-
lateral view.
WANG: EAST ASIA COELOTINE SPIDERS
FiGurE 43. Draconarius nanyuensis (Peng and Yin). A. Epigynum. B. Vulva.
607
608 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGureE 44. Draconarius neixiangensis (Hu, Wang and Wang). A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D.
Pedipalpus, retrolateral view.
basal haematodocha
conductor dorsal apophysis
tegulum
x ye /
subtegulum i
aematodocha
J > basal
‘
conductor lamella °°
_*
Figure 45. Draconarius nudulus Wang, sp. nov. A. Pedipalpus, ventral view. B. Pedipalpus, retrolateral view.
(Haematodochae slightly expanded).
WANG: EAST ASIA COELOTINE SPIDERS 609
WH (1%
q conductor dorsal AES
FIGURE 46. Draconarius ornatus (Wang et al.). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view. B.
Pedipalpus, retrolateral view.
FIGURE 47. Draconarius parabrunneus Wang, sp. nov. A. Epigynum. B. Vulva.
spermathecal head
epigynal hood
diverticulum
FiGuRE 48. Draconarius paraterebratus Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D.
Pedipalpus, retrolateral view.
610 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGURE 49. Draconarius patellabifidus Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D.
Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 611
ws conductor dorsal apophysis
=e SS
FicurE 50. Draconarius penicillatus (Wang et al.). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D.
Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
FiGURE 51. Draconarius potanini (Schenkel). A. Epigynum. B. Vulva.
612 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 53. Draconarius pseudocapitulatus Wang, sp. nov. A. Epigynum. B. Vulva.
epigynal tooth copulatory duct
a spetmathecal stalk
atrium fertilization duct -«‘SPet™mathecal base
FiGuRE 54. Draconarius pseudowuermlii Wang, sp. nov. A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 613
FIGURE 56. Draconarius rotundus Wang, sp. nov. A. Epigynum. B. Vulva.
614 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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spermathecal head copulatory duct
FiGurE 57. Draconarius rufulus (Wang et al.). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus,
ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS
FiGuRE 59. Draconarius simplicidens Wang, sp. nov. A. Epigynum. B. Vulva.
616 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 60. Draconarius singulatus (Wang et al.). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus,
ventral view. E. Pedipalpus, retrolateral view.
617
WANG: EAST ASIA COELOTINE SPIDERS
eri (Brignoli). A. Epigynum. B. Vulva.
sStemn
FIGURE 61. Draconarius
rius Striolatus (Wang et al.). A. Epigynum. B. Vulva.
FIGURE 62. Dracona
618 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 63. Draconarius terebratus (Peng and Wang). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS
619
FIGURE 64. Draconarius tibetensis Wang, sp. nov. A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view.
C. Pedipalpus, retrolateral view.
620 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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See esas
s(x
= ee an:
Sees ee eu
=o5 por f Bett
. Se Se _ Bs
> Bec,
< Xe ¥ \ a:
ey WS} I
FIGURE 65. Draconarius uncinatus (Wang et al.). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view.
C. Pedipalpus, retrolateral view.
FIGURE 66. Draconarius wenzhouensis (Chen). A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 621
FIGURE 67. Draconarius wudangensis (Chen and Zhao). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D.
Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
622 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 69. Draconarius yichengensis Wang, sp. nov. A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS
623
FIGURE 70. Femoracoelotes latus (Wang, Tso and Wu). A.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
624 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 71. Femoracoelotes platnicki (Wang and Ono). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, retrolateral view. E. Pedipalpus, ventral view.
WANG: EAST ASIA COELOTINE SPIDERS
FIGURE 72. Leptocoelotes edentulus (Wang and Ono). A. Epigynum. B. Vulva.
625
626 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 73. Leptocoelotes pseudoluniformis (Zhang, Peng and Kim). A, C. Epigyna, showing variation. B, D. Vulvae,
showing variation. E. Pedipalpus, prolateral view. F. Pedipalpus, ventral view. G. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 627
Ficure 74. Longicoelotes karschi Wang. A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus, ventral
view. E. Pedipalpus, retrolateral view.
628 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FicuRE 75. Platocoelotes impletus (Peng and Wang). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 629
FIGURE 76. Platocoelotes icohamatoides (Peng and Wang). A. Epigynum. B. Vulva.
630 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SSS
SSS QA . Ss ate ~ 5.
(dog
“conductor
Fh
FIGURE 77. Platocoelotes kailiensis Wang, sp. nov. A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 631
FIGURE 79. Spiricoelotes pseudozonatus Wang, sp. nov. A. Epigynum. B. Vulva.
632 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 80. Spiricoelotes zonatus (Peng and Wang). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 633
FiGuRE 81. Tegecoelotes secundus (Paik). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus, ven-
tral view. E, F. Pedipalpus, retrolateral view.
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FIGURE 82. Tegecoelotes corasides (B6senberg and Strand). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view.
D. Pedipalpus, ventral view. E, F. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 635
FIGURE 84. Tegecoelotes muscicapus (Bésenberg and Strand). A. Pedipalpus, prolateral view. B. Pedipalpus, ventral
view. C. Pedipalpus, retrolateral view.
636 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 86. Tonsilla truculenta Wang and Yin, female holotype, male allotype, from Tianzishan, Sangzhi, Hunan, China.
A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus, ventral view. E. Pedipalpus, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 637
FIGURE 87. Tonsilla truculenta Wang and Yin, variation one, male and female paratypes from Tianzishan, Sangzhi,
Hunan, China, the same locality as holotype female. A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D. Pedipalpus,
retrolateral view.
FIGURE 88. Tonsilla truculenta Wang and Yin, variation two, female paratype from Tianzishan, Sangzhi, Hunan, China,
the same locality as holotype female. A. Epigynum. B. Vulva.
638 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FicurE 89. Tonsilla truculenta Wang and Yin, variation three, females and male from Guiyang, Guizhou, China.
A. Epigynum. B. Vulva. C. Pedipalpus, ventral view. D, E. Pedipalpi, retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 639
FiGurE 90. Tonsilla truculenta Wang and Yin, variation four, male from Guiyang, Guizhou, China. A. Pedipalpus, ven-
tral view. B. Pedipalpus, retrolateral view.
640 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 91. Tonsilla truculenta Wang and Yin, variation five, female paratype from Meitan, Guizhou, China. A.
Epigynum. B. Vulva.
A
FIGURE 92. Tonsilla lyratus (Wang et al.). A. Epigynum. B. Vulva.
A
FIGURE 93. Tonsilla tautispinus (Wang et al.). A. Epigynum. B. Vulva.
WANG: EAST ASIA COELOTINE SPIDERS 641
FiGuRE 94. Tonsilla variegatus (Wang et al.). A. Epigynum. B. Vulva. C. Pedipalpus, prolateral view. D. Pedipalpus,
ventral view. E, F. Pedipalpi, retrolateral view.
642 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 95. Tonsilla makros Wang, sp. nov. A. Pedipalpus, prolateral view. B. Pedipalpus, ventral view. C. Pedipalpus,
retrolateral view.
WANG: EAST ASIA COELOTINE SPIDERS 643
FiGuRE 96. Habitus of Draconarius, dorsal views. A. Draconarius haopingensis, female. B. Draconarius jiangyongen-
sis, female. C. Draconarius ornatus, male. D. Draconarius molluscus, female. E. Draconarius parabrunneus, female.
F. Draconarius uncinatus, male. G. Draconarius terebratus, female. H. Draconarius terebratus, male. 1. Draconarius rufu-
lus, female.
644 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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WANG: EAST ASIA COELOTINE SPIDERS 645
FiGuRE 97. Habitus of Coelotinae, dorsal views. A. Draconarius acidentatus, female. B. Draconarius adligansus,
female. C. Draconarius baxiantaiensis, female. D. Draconarius calcariformis, female. E. Draconarius chaigiaoensis,
female. F. Draconarius shuangpaiensis (= Draconarius digitusiformis), male. G. Bifidocoelotes primus, female. H. Tonsilla
lyratus, female. I. Spiricoelotes zonatus, female.
646 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Bifidocoelotes
B bifida
@ primus
Map 1. East Asia, showing distribution of Bifidocoelotes species.
Coronilla eae HD ae
m gemata Se
@ libo poy
A Sigillata aa ee
© subsigillata rae
& magnshan ; “acs = en
Map 2. East Asia, showing distribution of Coronilla species.
WANG: EAST ASIA COELOTINE SPIDERS 647
Draconarius
mg acidentatus
@ absentis
A adligansus
® al/tissimus
—_ =
Map 3. East Asia, showing distribution of Draconarius absentis, D. acidentatus, D. adligansus, and D.
altissimus.
Draconarius
gw arcuatus
@ amygdaliformis
A argenteus
® agrestis
Map 4. East Asia, showing distribution of Draconarius agrestis, D. amygdaliformis, D. arcuatus, and D.
argenteus.
648 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconarius
B aspinatus
@ baronii
é baxiantaiensis
© brunneus
Map 5. East Asia, showing distribution of Draconarius aspinatus, D. baronii, D. baxiantaiensis, and D.
brunneus.
Draconarius
@ bituberculatus
@ calcariformis
A capitulatus
© chaigiaoensis
Map 6. East Asia, showing distribution of Draconarius bituberculatus, D. calcariformis, D. capitulatus, and
D. chaigiaoensis.
WANG: EAST ASIA COELOTINE SPIDERS 649
Draconarius
& colubrinus
@ carinatus
@ cheni
& coreanus
® curiosus
Draconarius
m davidi
@ denisi =
A digitusiformis —~
® disgregus
Map 8. East Asia, showing distribution of Draconarius davidi, D. denisi, D. digitusiformis, and D. disgre-
gus.
650 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconarius
@ dissitus
@ dubius
é everesti
® funiushanensis —_~
Map 9. East Asia, showing distribution of Draconarius dissitus, D. dubius, D. everesti, and D. funiushanen-
SIS.
Draconarius
m@ episomos
@ gurkha
A gyriniformis
© haopingensis
Map 10. East Asia, showing distribution of Draconarius episomos, D. gurkha, D. gyriniformis, and D.
haopingensis.
WANG: EAST ASIA COELOTINE SPIDERS 651
Draconarius
mw griswoldi
@ hangzhouensis
A himalayaensis
© huizhunesis
Map 11. East Asia, showing distribution of Draconarius griswoldi, D. hangzhouensis, D. himalayaensis, and
D. huizhunesis.
Draconarius
gw hui
@ incertus
A infulatus
@jiangyongensis
S Za LD
Map 12. East Asia, showing distribution of Draconarius hui, D. incertus, D. infulatus, and D. jiangyongen-
SiS.
652 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconarius
g /abiatus
@ /inzhiensis
A /inxiaensis
®/utulentus
Map 13. East Asia, showing distribution of Draconarius labiatus, D. linzhiensis, D. linxiaensis, and D. lutu-
lentus.
Draconarius
m Magniceps
@ molluscus
A nanyuensis
@neixiangensis
Map 14. East Asia, showing distribution of Draconarius magniceps, D. molluscus, D. nanyuensis, and D.
neixiangensis.
WANG: EAST ASIA COELOTINE SPIDERS
Draconarius
m nudulus
@ ornatus
& parabrunneus
@pervicax ~~
Map 15. East Asia, showing distribution of Draconarius nudulus, D. ornatus, D. parabrunneus, and D. per-
vicax.
Draconarius
Bite ra @ paraterebratus
=k @ penicillatus
A picta
© potanini
Map 16. East Asia, showing distribution of Draconarius paraterebratus, D. penicillatus, D. picta, and D.
potanini.
653
654 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
Draconanus
@ patellabifidus
@ gingzangensis
A guadratus
© rotundus
Map 17. East Asia, showing distribution of Draconarius patellabifidus, D. gingzangensis, D. quadratus, and
D. rotundus.
Draconarius
m pseudowuermlii
@ pseudobrunneus
A rufulus
© schenkeli
Map 18. East Asia, showing distribution of Draconarius pseudowuermlii, D. pseudobrunneus, D. rufulus,
and D. schenkeli.
WANG: EAST ASIA COELOTINE SPIDERS 655
Draconarius
m@ pseudocapitulatus
@ simplicidens
&é singulatus
© stemmleri
Map 19. East Asia, showing distribution of Draconarius pseudocapitulatus, D. simplicidens, D. singulatus,
and D. stemmleri.
Draconarius
m streptus
@ striolatus
A strophadatus
© subtitanu
nen
Map 20. East Asia, showing distribution of Draconarius streptus, D. striolatus, D. strophadatus, and D. sub-
titanus.
656 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Draconanus
m syzygiatus
@ terebratus
A tibetensis
© tryblionatus ~
Map 21. East Asia, showing distribution of Draconarius syzygiatus, D. terebratus, D. tibetensis, and D. try-
blionatus.
ee
Ss
@ trifasciatus
@ uncinatus
A venustus
———
© wenzhouensis |
Map 22. East Asia, showing distribution of Draconarius trifasciatus, D. uncinatus, D. venustus, and D. wen-
zhouensis.
WANG: EAST ASIA COELOTINE SPIDERS 657
Draconarius
m Wwudangensis
@ wuermiii LS
A yadongensis _
® yichengensis
Map 23. East Asia, showing distribution of Draconarius wudangensis, D. wuermlii, D. yadongensis, and D.
yichengensis.
Map 24. East Asia, showing distribution of Draconarius yosiianus.
658 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Femoracoelotes
g /atus
e platnicki
eo so
Leptocoelotes
mg edentulus
@ pseudoluniformis
Map 26. East Asia, showing distribution of Leptocoelotes edentulus and L. pseudoluniformis.
WANG: EAST ASIA COELOTINE SPIDERS
Longicoelotes
mkulinganus
e senkakuensis ~
& karschi
Platocoelotes
gw /mpletus
@ icohamatoides
é icohamatus
® kailiensis
0 lichuanensis
Map 28. East Asia, showing distribution of Platocoelotes impletus, P. icohamatoides, P. icohamatus, P.
kailiensis, and P. lichuanensis.
659
660 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Spiricoelotes
m pseudozonatus’
@ urumensis
& zonatus-
Tegecoelotes
m secunda
@ corasides
& jgnotus
© michikoae
© muscicapa
Map 30. East Asia, showing distribution of Tegecoelotes secundus, T. corasides, T. ignotus, T. michikoae, and
T. muscicapus. (the distribution site data are incomplete)
WANG: EAST ASIA COELOTINE SPIDERS
= Jonsilla eburniformis A Tonsilla imitata
© Tonsilla truculenta ® Tonsilla truculenta (variation 1)
© Tonsilla truculenta (variation 2) © Tonsilla truculenta (variation 3)
= Jonsilla truculenta (variation 4) © Tonsilla truculenta (variation 5)
Map 31. East Asia, showing distribution of Tonsilla eburniformis, T. imitata, and T: truculenta.
661
662 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 26
ee Tonsilla
@ /yratus
@ tautispinus
A variegatus
© makros
Map 33. East Asia, showing distribution of Draconarius.
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27, pp. 663-686, 10 figs., 1 table, Appendix November 14, 2003
Fact, Theory and Tradition in the Study of Molluscan Origins
David R. Lindberg!” and Michael T. Ghiselin?
! Department of Integrative Biology and Museum of Paleontology, University of California,
Berkeley, CA, USA 94720-4780, email: drl@uclink.berkeley.edu; * Center for the History and Philosophy
of Science and Department of Invertebrate Zoology and Geology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA, USA 94118, email: mghiselin@calacademy.org.
Discussions of phylogenetic relationships and origins often use figures called ‘‘arche-
types,” or ‘treconstructed common ancestors.” Here we discuss one such creature,
the hypothetical ancestral mollusc or HAM. HAM first appeared 150 years ago as
T.H. Huxley’s archetypal mollusc and has speciated often since then. Radiations
have occurred within both fossil and Recent taxa (from the paleontological and
neontological literature, respectively). Eight species have appeared in the last 30
years alone and at least six species remain extant today.
We performed both phenetic and cladistic analyses of the character states present
in these figures. Our best approximation of the phylogeny of HAM (based on known
ancestor-descendant relationships and stratigraphy) requires 53 more steps than the
most parsimonious tree found by cladistic analysis. Phenetic trees based on neighbor
joining and UPGMA analyses require two and twelve more steps, respectively than
the most parsimonious trees. The evolution of HAM exhibits all the typical process-
es and developmental heterochronies thought to encompass organic morphological
evolution, and both phenetic analysis and cladistic analyses have problems relating
paedomorphic taxa.
HAM has not aided evolutionary biologists or paleontologists in solving problems,
but it has often had the opposite effect, by requiring that theories be treated within
its framework. Moreover, real data have ended up being ‘“‘tested”’ against a hypothet-
ical anatomy to determine whether a hypothesis should be accepted or rejected. It
has been argued that HAM serves a valuable role as a pedagogical teaching aid.
Unfortunately, these imaginary animals do not come clearly labeled with warnings
about the harm that they might do if mistaken for real organisms.
Key words: Mollusca; hypothetical ancestors; phylogeny; archetypes; typology; conceptual evolution.
According to the current evolutionary ontology, species, lineages, and many other things are
individuals, not classes (Ghiselin 1974, 1997; Hull 1976). This suggests that at least some of those
“other things” might be studied from an evolutionary point of view. Among the candidates have
been sport cars (Rowland 1968), literary genres (Ghiselin 1980), cladists (Carpenter 1987), and sci-
entific theories (Hull 1988). The list could be greatly expanded. One intriguing possibility is line-
ages of diagrams in the scientific literature. Griesemer and Wimsatt (1989) examine the diagrams
that illustrate the Weismannian concept of the continuity of the germ plasm and discuss the study
of such diagrams in considerable depth. As they point out, such diagrams can be dated precisely,
3 Research Associate, Department of Invertebrate Zoology and Geology, California Academy of Sciences.
663
664 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
are a very convenient focus of attention, and the analogy between a given diagram and an organ-
ism that forms part of a lineage is captivating.
Here we consider some diagrams that depict something even more strongly analogous (and
perhaps in a certain sense homologous) to organisms: pictures of hypothetical ancestral molluscs
that adorn (one might prefer to say disfigure) the pages of the scientific literature. Our study dif-
fers from that of Griesemer and Wimsatt by incorporating phenetic and cladistic analyses to esti-
mate the relationships and character transformations of the images. Furthermore, we direct our
commentary primarily to comparative biologists and paleobiologists like ourselves, though we
hope that historians and philosophers of science will find it useful as well.
Discussions of phylogenetic problems often use figures called “archetypes,” “Baupldne,” or
“reconstructed common ancestors.” There is some disagreement as to what such diagrams are sup-
posed to mean. Idealistic morphologists, such as Remane (1952) have asserted that they are just
diagrams of the relative position of parts, designed, perhaps, to look like animals. Others, such as
Beklemischew (1969), have claimed that, if done properly, they really do represent the common
ancestor. From the point of view of taxonomy, we might consider them “pictorial diagnoses”—
summaries of the features common to all the organisms in a taxon. From the point of view of phy-
logenetics, we might consider them hypotheses about a common ancestor — illustrating what was
inferred to be present in the last common ancestor of the group. These interpretations are not mutu-
ally exclusive.
Systematists often treat hypotheses about common ancestors as results, rather than as starting
points, of investigations. To do so makes sense only in terms of a naive inductionism that does no
justice to the subtleties of scientific reasoning. Hypotheses are conjectures that are tested by refer-
ence to evidence, perhaps by refuting them, as suggested by the title of a well-known book by
Popper (1962). Whether one is a “refutationist” (who would deny that hypotheses can be verified)
or not, a hypothesis remains a hypothesis irrespective of whether one has gathered any evidence
that might lead one to prefer it or some alternative.
Hypotheses are tested by means of experiments, or by comparative methods that are of the
same basic logical form as experiments. When a hypothesis explains a variety of empirical data and
withstands serious attempts at refutation, it is preferable to one that explains less or generates false
predictions about the material universe. With respect to hypothetical ancestral models, this means
that at the very least they should have the properties that biologists find among living organisms in
general. But the mere plausibility of such a model is no reason to prefer it to some alternative that
is equally plausible, or perhaps even more so.
Nonetheless, hypotheses often remain viable in spite of contradictory evidence. There may be
some conditions under which it is scientifically legitimate not to falsify a particular hypothesis
even when it seems to have been refuted. For example, the test may refute, not the hypothesis itself,
but another premise in the system, for example, a “fact” that turns out to be an inaccurate measure-
ment. However, when the hypothesis is simply taken for granted, and contradictory data are
ignored or explained away, something is wrong. In this case we are dealing with something that
functions not as a hypothesis, but as an assumption being treated as if it were fact. Non-algorith-
mic phylogenetic hypotheses are often treated this way, but so are many others. The reasons for this
are not to be sought in pure logic. Scientists are not alone in accepting what everybody around them
takes for granted. They believe what they read in textbooks and what they were told by their teach-
ers. What they accept becomes an integral part of a body of knowledge, in which fact and conjec-
ture are not clearly distinguished. The data then become selectively gathered in a way that tends to
“confirm” the assumptions. If unaware of the premises, scientists may be unable to recognize the
circularity of their own reasoning. Thus, we get covert assumptions that are better called “tradi-
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 665
tions” than “hypotheses,” but that are treated as “facts” by their proponents. Only by going back to
fundamentals, identifying the premises, and distinguishing fact from theory, can we hope to get
outside the circle. The initial premises must be conjectural and open to revision. The notion that
one can start out with unbiased data is a preposterous myth.
Traditions, like species, are individuals: they originate, vary, compete with one another,
evolve, and, ultimately, become extinct. To come to grips with them, we must deal with them as
historical entities, see where they came from, and what forces have maintained them in their envi-
ronments. Herein we discuss one such creature, Huxley’s Archetypical Mollusc — HAM for short.
Kept in an artificial, polemic environment, it has flourished and persisted in spite of its teratologi-
cal character, an animal that never existed except in the fancy of theoreticians.
MATERIALS AND METHODS
TAXA STUDIED.— Our study lineage dates from Huxley (1853) who referred to his special cre-
ation as the "archetypal molluscous form" and cautioned that he was not proposing any idealized
form. Instead, Huxley viewed his archetype as a starting point on which the "the known laws of
development” might act to modify it into “the different secondary types.” Thirty years later
Lankester (1883) referred to his modification of Huxley's diagram as the "schematic mollusc" in
the figure caption. However, within the text Lankester allowed for the possibility that the schemat-
ic mollusc might, in fact, represent the “original Mollusc or archi-Mollusc (more correctly
Archimalakion).” For consistency we here select Huxley's use of the phrase “‘archetypal mollus-
cous form” as the first designation, and because of the lack of rules of nomenclature for common
names, and as first revisers of HAM nomenclature (neither of which matters), transmute this to
“Huxley's Archetypal Mollusc” or HAM, and designate it as the type species and the common
ancestor of all taxa placed in the imaginary genus Hamus.
Specimens collected in academic libraries provided clear evidence that HAM has speciated
often since 1853. At least eight species have appeared in the last 30 years alone and more than six
species remain extant today. These species include both fossil and Recent taxa (from the paleonto-
logical and neontological literature respectively). Twenty-two taxa were included in the initial
analyses; the outgroup taxon and 21 ingroup taxa (Table | and Appendix). These taxa were chosen
because they represented both neontological and paleontological lineages, and are mostly found in
textbooks.
Six additional “Cambrian” species were included in a second analysis. These species have a
first occurrence in Pojeta and Runnegar (1976: Figs. 9a-f), and have had a very important role in
the discussion of monoplacophoran and gastropod relationships (e.g., Peel 1991; Geyer 1986;
Parkhaev 2001). We included them here in a second analysis because fossils can often be extreme-
ly important in phylogenetic reconstruction (Donoghue et al. 1989).
EXCLUDED TAXA. — We restrict our analysis of taxa primarily to the Anglo-American subclade
Hamus s.s. Other taxa, belonging to separate intellectual traditions (i.e., lineages) have evolved in
relative isolation to the Anglo-American fauna. For example, many of the German language taxa
are Clearly distinct, the language barrier inducing a certain amount of endemism and reduced
opportunities for outbreeding and hybridization although there is some evidence of limited out-
crossing on the continent. For example, Haszprunar (1992) published a HAM in an Italian journal
and Salvini-Plawen and Steiner (1995) published in an English book (we include Hamus salvinis-
teinerorum in our analysis because of its prominent Anglo habitat). Also see remarks on the rela-
tionships of Hamus pelseneeri, Hamus (Jambonus) portmanni and Hamus (Schinkenus) naefi
below.
Other taxa such as the peculiar Protohamus verrilli developed by Verrill (1896) and
666 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
Protohamus yongi (Yonge 1960) have been excluded from our analysis. Verrill's species was a
larva or “veliger-like form”; an enviable hopeful monster, evidently produced through complete
progenetic loss of all adult structures. P. yongi is similar to Verrill's creation, although Yonge's later
special creations are clearly referable to the genus Hamus.
OUTGROUP AND CHARACTER SELECTION.— The data available for the study of the evolution of
Hamus are unique. We possess an almost 2000
perfect fossil record for the taxon, pre-
1980
served in dated textbooks and other publi-
cations (Fig. 1; Table 1). We also have the 1960 5
actual common ancestor in its natural 4940
environment. The characters are limited to §&
: > 1920
the anatomical features that each author
penned into their creature, and the charac- 1900 +
ter transformations fixed by our possession 1880 -
of most of the actual phylogeny of the
group, often indicated by such expressions chy
as “After. . .”, or “Redrawn from...” in the
caption of the figure. In tracing the various Taxon Number
derived forms from their known origin, we FicurE 1. Stratigraphic distribution of Hamus species treated
can readily observe the character transfor- herein. Taxa 10 and 18—22 are currently extant. Taxon numbers are
mations, evolutionary trends and diver- RUSS RS EL
gences. Moreover, we have an excellent opportunity to test phylogenetic hypotheses (e.g.,
phenograms and cladograms) with the actual history of the group.
HAM did not arise without antecedents. Traces of its early beginnings can be found in the writ-
ings of von Baer (1828:pl. 3, fig. 12). However, as a clearly recognizable entity (i.e., an individ-
ual), from which the others were derived, it takes its origin from a drawing by T.H. Huxley (1853)
(Fig. 2). Therefore, Hamus huxleyi is the consummate sister taxon and outgroup.
The ancestral condition exhibited by Hamus huxleyi provides the plesiomorphic states for
characters used in our analysis (Fig. 2;
Appendix). These character states include
a body with clearly differentiated head
and foot, cephalic and optic tentacles, and
a dorsal visceral mass covered by a man-
tle. Surprisingly, a shell is absent. The vis-
cera consist of a straight digestive tract
with a style sac, a heart with an anterior
aorta, and paired kidneys. Gills are present FIGURE 2. Hamus huxleyi. T.H. Huxley’s Archetype of the
: Rote = : Cephalous Mollusca. Redrawn from Huxley (1853).
on the posterio-dorsal surface of the ani-
mal. It is important to note that in the ancestral condition the gills are not located in a posterior
mantle cavity. The H. huxleyi nervous system consists of four major pairs of ganglia (cerebral,
pleural, pedal, and buccal). Obviously Huxley had a rather snail-like ancestor in mind — a sort of
untorted limpet without a shell. It stands to reason that when one thinks typologically one will asso-
ciate “the mollusc” with the most familiar examples, rather than with creatures that are more exot-
ic such as chitons or scaphopods.
ANALYsIS.— The character matrix and analysis (see Appendix) describes the 36 characters
used in our analyses and their states. We performed both phenetic and cladistic analyses of the data
using PAUP* Ver. 4.0610 (Swofford 1998). The UPGMA and neighbor joining options in PAUP*
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY
TABLE |. Data for Hamus species analyzed here. Citations for figured specimens consists of Author(s) + First
columns. Data present below are also graphically represented in Figure | (First and Last occurrences) and
Figure 8a (Duration).
667
First
Duration Paleo Textbooks Insular
Taxon Taxon Author(s) Last
No. (to 2000)
1 H. huxleyi Huxley 1853 wl883 30 Fy
2 H. lankesteri Lankester 1883 1906 23 |
3 H. nicholsoni Nicholson & 1889 1923 34 \ V
Lydekker,
4 H. pelseneeri Pelseneer 1906 1920 14 V V
5 H. swinnertoni Swinnerton 1923 1943 20 V V
6 H. borrapottsorum Borradaile & Potts 1932 1967 39 V V
7 H. moolafisherorum Moore, Lalicker & 1952 1965 13 V
Fisher
8 H. mortoni Morton 1958 1967 9 V
9-H. eastoni Easton 1960 1967 7 V
10 _-H.. barnesi Barnes 1963 — ai) V
11 H. mortonyongorum Morton & Yonge 1964 1983 19 V
12H. hickmani Hickman 1967 1978 ll V
13 H. meglitschi Meglitsch 1967 199] 24 V
14H. russellhunteri Russell-Hunter 1968 1979 11 V
15 H. staseki Stasek 1972 1982 10
16 H. yongethompsonorum Yonge & 1976 1982 6 V
Thompson
17 H. barthbrosorum Barth & Broshears 1982 1992 10 V
18 H. bossi Boss 1982 — 18
19H. seedi Seed 1983 — iy V
20. _-H. barcalolivorum Barnes, Calow & 1988 = 12 V V
Olive
21 H. ravenjohnsonorum Raven & Johnson 1992 — 8 V
22 H. salvinisteinerorum Salvini-Plawen & 1996 —- 4
Steiner
were used to calculate phenetic trees based on distance matrix of mean character differences. In
both phenetic analyses ties were broken randomly. The data matrix was also subjected to heuristic
searches under maximum parsimony with H. huxleyi serving as the outgroup. All characters were
equally weighted and unordered, and were assumed to show accelerated character transformation.
Strict and majority rule (50%) consensus trees were calculated.
RESULTS
The single tree from the UPGMA analysis is presented in Fig. 3a; it contains two distinct
groupings. The first group is made up of the early HAM taxa + paedomorphic taxa + flatworm-like
668
0.1
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
H. huxleyi
H.staseki
H.lankesteri
H.nicholsoni
H.moolafisherorum
H.swinnertoni
H.eastoni
H.russellhunteri
H.bossi
H.ravenjohnsonorum
H.salvinisteinerorum
H.barcalolivorum
H. pelseneeri
H.borrapottsorum
H.meglitschi
H.hickmani
H.barthbrosorum
H.mortoni
H.mortonyongorum
H.yongethompsonorum
H.barnesi
H.seedi
H. huxleyi
H.lankesteri
H.nicholsoni
H.borrapottsorum
0.1
FiGuRE 3. Phenetic trees from analysis of distance matrix of mean char-
acter differences. a. UPGMA and b. neighbor joining analysis. (See
Appendix for data matrix.)
H.pelseneeri
H.staseki
H.bossi
H.moolafisherorum
H.swinnertoni
H.eastoni
H.russellhunteri
H.ravenjohnsonorum
H.salvinisteinerorum
H.barcalolivorum
H.mortonyongorum
H.mortoni
UW H. yongethompsonorum
H.barnesi
H.seedi
H.meglitschi
H.hickmani
H.barthbrosorum
Volume 54, No. 27
taxa. Although the paedomorphs +
flatworm-like taxa are clearly
delimited in the analysis, the dis-
junct stratigraphic record of these
taxa suggests that this cluster is
better attributed to convergence
rather than common ancestry. The
second cluster consists primarily
of textbook HAMs + the insular
(i.e., British) HAMs. Unlike the
first cluster, the textbook and insu-
lar HAMs do not form unique
groups, but instead are mixed
within the cluster. The topology of
the UPGMA tree requires 108
steps.
The single tree from the
neighbor joining analysis is pre-
sented in Fig. 3b. In the neighbor
joining analysis, the flatworm-like
taxa group with the textbook and
insular HAMs with the paedo-
morph group as its sister taxon.
Hamus_ lankesteri and Hamus
nicholsoni are outside of these
three groups in both phenetic
analyses. The topology of the
neighbor joining tree requires 98
steps.
Maximum parsimony analysis
found 76 trees with 96 character
steps. Strict and majority rule con-
sensus trees for these trees are pre-
sented in Fig. 4. In the strict con-
sensus tree, four subclades are
present (Fig. 4a). These are: (1)
Hamus mortoni and Hamus
yongethompsonorum, (2) Hamus
pelseneeri and the flat worm-like
HAMs, (3) the paedomorph group,
and (4) HAMs from four American
invertebrate zoology textbooks
and the single insular species
Hamus seedi. The majority rule
consensus tree (Fig. 4b) is virtually identical to the neighbor joining solution; they differ primari-
ly in the placement of Hamus mortonyongorum and Hamus borrapottsorum. Half of the resolved
branches appear in 80% or more of the most parsimonious trees (Fig. 4b).
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 669
H. huxleyi
a H.lankesteri
H.nicholsoni
H.borrapottsorum
H.moolafisherorum
H.mortonyongorum
H.barcalolivorum
H. pelseneeri
H.staseki
H.bossi
H.ravenjohnsonorum
H.salvinisteinerorum
H.swinnertoni
H.eastoni
H.russellhunteri
H.mortoni
H.yongethompsonorum
H.barnesi
H.hickmani
H.meglitschi
H.barthbrosorum
H.seedi
H. huxleyi
b H.lankesteri
H.nicholsoni
H.borrapottsorum
H.pelseneeri
H.staseki
H.bossi
H.ravenjohnsonorum
H.salvinisteinerorum
H.moolafisherorum
H.swinnertoni
H.eastoni
H.russellhunteri
H.barcalolivorum
H.mortonyongorum
H.mortoni
H.yongethompsonorum
H.hickmant
H.meglitschi
H.barthbrosorum
H.barnesi
H.seedi
FicurE 4. Consensus cladograms of 76 most parsimonious trees found by
PAUP* analysies of the 22 HAM taxa listed in Table 1. CI and RI for randomly
selected tree (No. 127) = 0.4725 and 0.6000, respectively. a. Strict and b. majority
rule consensus trees. (See Appendix for data matrix.)
In the final cladistic
analysis, the “Cambrian”
species complex Hamus
pojetarunnegarorum, com-
prising six taxa, was added
to the data matrix. Hamus
pojetarunnegarorum taxa
are well known in the
Cambrian literature (e.g.,
Pojeta and Runnegar 1976)
Where they are typically
used to differentiate be-
tween untorted and torted
molluscs. Because of their
association with some of the
oldest molluscan fossils,
they have some of the high-
est number of unknown
character states in the analy-
sis, being surpassed only by
some of the paedomorphic
taxa (see Appendix). The
addition of these taxa to the
data matrix produced 4875
trees of 1034 character
steps. In the strict consensus
tree, the addition of these
taxa produces a large unre-
solved polytomy at the base
of the tree while the only
remaining subclades are
Hamus mortoni and Hamus
yongethompsonorum — and
the HAMs_ from _ four
American invertebrate zool-
ogy textbooks + the single
insular species Hamus seedi
(Fig. 5a). In the majority
rule consensus tree (Fig.
5b), the complex is situated
in a subclade consisting pri-
marily of paedomorphs with
the exception of the flat-
worm-like Hamus raven-
jJohnsonorum with which H.
pojetarunnegarorum taxon groups. The remaining members of the complex form their sister taxon.
The overall effect of the addition of fossils to our analysis is to intermingle members of the flat-
H. huxleyi
H.lankesteri
H.nicholsoni
H.pelseneeri
H.swinnertoni
H.borrapottsorum
H.moolafisherorum
H.eastoni
H.mortonyongorum
H.russellhunteri
H.staseki
H.bossi
H.barcalolivorum
H.ravenjohnsonorum
H.salvinisteinerorum
Taxona
Taxon b
Taxon c
Taxon d
Taxon e
Taxon f
H.mortoni
H.yongethompsonorum
H.barnesi
H.hickmani
H.meglitschi
H.barthbrosorum
H.seedi
H. huxleyi
H.moolafisherorum
H. borrapottsorum
H.lankesteri
H.nicholsoni
H.pelseneeri
H.staseki
H.bossi
H.salvinisteinerorum
H.swinnertoni
H.eastoni
H.russellhunteri
Taxon a
Taxon b
Taxonc
Taxon d
Taxone
H.ravenjohnsonorum
Taxon f
H.barcalolivorum
H.mortonyongorum
H.mortoni
H.yongethompsonorum
H.hickmani
H.meglitschi
H.barthbrosorum
H.barnesi
H.seedi
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
worm-like and paedomorph
groups and reduce overall resolu-
tion of these groups. In contrast,
occurrence of the subclade con-
sisting of Hamus mortonyongo-
rum, Hamus _ barcalolivorum,
Hamus mortoni, Hamus yongeth-
ompsonorum, the four American
invertebrate zoology textbook
HAMs and Hamus seedi increases
from 68% to 97% at three nodes
because of the addition of the fos-
sils (cf. Figs. 4b and 5b).
Our best approximation of the
actual phylogeny of HAM based
on known ancestor-descent rela-
tionships and stratigraphy (Fig. 6)
requires 53 more steps than the
most parsimonious trees. Three
distinctive clades are present in
this phylogeny (Fig. 6):
|. Lankester group.— an early
group (1883) of mostly fossil species
tracing their ancestry to Hamus
lankesteri. A single pair of paedo-
morphs represents the most “derived”
taxa within this clade.
2. Morton group.— a large, comb-
like group whose members trace their
ancestry to Hamus mortoni (1958).
This group consists of intermingled
insular and textbook Hamus species.
A second pair of paedomorphs is nest-
ed within this clade.
3. Stasek group.— the sister taxon of
the Morton group, and although it
shares the same minimum age of
divergence (1958) with the Morton
group, the earliest known members
date from 1972 (Hamus staseki). Half
of the extant Hamus species are mem-
bers of this group. The continental
taxon H. salvinisteinerorum is arbi-
trarily placed in this clade because of
its recurrent sister taxon relationship
with H. ravenjohnsonorum in all
analyses with the exception of the
addition of the Cambrian taxa.
FiGuRE 5. Consensus cladograms of 4875 most parsimonious trees found
by PAUP* analysies of the 22 HAM taxa listed in Table 1 plus six taxa of the
“Cambrian” species complex Hamus pojetarunnegarorum. Cl and RI for ran-
domly selected tree (No. 356) = 0.4757 and 0.6932, respectively. a. Strict and
b. majority rule consensus trees. (See Appendix for data matrix.)
Two of these groups were
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 671
H.huxleyi present to varying degrees in our
Pankosteri « amnalyses (cf. Figs. 3-6). Com-
1883 5
hi roetiorh © ponents of the Morton group are
ji ; 2 present in the UPGMA and
Erewinnerton g neighbor joining trees, and in
ae 1923 |__ H.eastoni 5S both the strict and majority rule
H.pelseneeri consensus cladograms. The
H.borrapottsorum Stasek group was represented by
LENO SHEFOrUR, the flatworm-like taxa in the
7G06 Ee UPGMA and neighbor joining
.stase oe
1972 . a analyses, and in the majority rule
H.bossi © consensus cladograms. In the
1952 H.ravenjohnsonorum | % UPGMA analysis the Stasek
H.salvinisteinerorum ® group was the sister taxon of the
Marion paedomorph group (Fig. 3a),
ry ane while in the majority rule con-
sensus cladogram, the neighbor
1964 1968 [— H.russellhunteri joining tree, and the true phy-
1958 H.barcalolivorum ca logeny, this group is the sister
o geny group
H.yongethompsonorum © taxon of the Morton group (cf.
¢
H.barthbrosorum 5 Figs. 3b, 4b, and 6).
= ee 99
1963 H.mortonyongorum ithe Hacdomorp f erOuP
which is present in some form in
ised! all of our analysis is polyphylet-
H.barnesi ic and represents two separate
H.meglitschi events, one in the Lankester
FicureE 6. Best approximation of the actual relationships of Hamus species group and the other within the
based on known ancestor-descent relationships and stratigraphy. Dates at Morton group. We regard these
nodes indicate minimum divergence times for sister taxa. Consistency index =
0.2986, retention index = 0.1583. MEDS) gs paedomorphs because of
their degenerated morphology,
and their anatomies have converged on each other as well as with the fossil taxa (Fig. 5b).
However, there is a 45 year difference in the time of origination of the first and most recent of these
groups and any suggested relationship is likely to be spurious. The universal occurrence of this
group in our analyses shows that whether real organisms or cartoons, paedomorphs are difficult
taxa to relate in phylogenetic reconstructions.
STRATIGRAPHY.— Fossil HAMs appear earlier in the record then the majority of zoological
ones (Fig. 1, Table 1). However, not one fossil HAM has originated in a textbook environment
since 1964. In contrast, 76% of zoological HAMs have originated since 1960 and all six of the
extant HAMs (7.e., texts and monographs not out of print) are zoological species (Table 1). Before
1960, the ratio between fossil and zoological Hamus species was about 1:1 (Table 1).
Following Norell and Novacek (1992) we compared the known stratigraphic ranges with
divergence patterns based on our cladistic and phenetic analyses. Testing the fossil record against
cladistic phylogenies typically determines how complete the record probably is, and therefore how
useful the taxon might be for stratigraphic purposes (Padian et al. 1994). However in our unique
situation, the stratigraphy is certain and therefore can be used to evaluate the reconstructed diver-
gence patterns (Fig. 7).
When the number of branch nodes between Hamus huxleyi and the taxa on the true tree are
672
os ct}
on
Number of nodes from origin
ort NN WO fH TH DN OO OO
lo)
fo)
\
\
(e)
oO
e
e
3
\
\
e e e -# ee
oe
e yo 8
i
4 fe) ° fo) ~-00®e fo) 9.0
By i
4 @o & 008-5 0 fe)
8 >SO O
= OmOwe anne ne ° fe)
ma
Number of nodes from origin
o-r NW fF IA DN OO OO
Rank of appearance
FIGURE 7. Scatterplots of the relationship between rank
of appearance (as measured by absolute age) and the num-
ber of branch nodes between origin and taxon on tree. a.
Results plotted from cladistic analyses. Solid circles from
strict consensus cladogram (72 = 0.1312, slope = 0.0335);
open circles from majority rule consensus cladogram (r2 =
0.5710, slope = 0.3571); b. Results plotted from phenetic
analyses. Solid circles from neighbor joining tree (r2 =
0.5900, slope = 0.3920); open circles from UPGMA
phenogram (r2 = 0.2299, slope = 0.1400).
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
plotted on the rank of appearance of the taxa,
the resulting 7° of the regression equals 0.6746
and the slope equals 0.4223. This is not signif-
icantly different from the majority rule consen-
sus tree (Fig. 7a) or the phenetic UPGMA tree
(Fig. 7b) (pairwise t-test, p > 0.05). However, it
is significantly different (p = 0.0012) from the
neighbor joining tree regression (Fig. 7b). The
strict consensus tree has virtually no strati-
graphic signal with a slope of 0.0345 (Fig. 7a).
Origination rates remained below 0.15
species/year for the first 80 years of Hammian
history, however between 1960 and 1980 origi-
nation rates skyrocketed to over 0.40 species/
year and have since begun to decline (Fig. 8a).
Extinction rates have typically been lower, but
follow a similar trend (Fig. 8a). The conse-
quence of the difference between these two
rates is a roughly exponential species diversity
curve through time (Fig. 8b). Although species
diversity was <2 for over 100 years, the sub-
stantially higher origination rates have main-
tained a relatively stable diversity of five or
more HAMs for the last 30 years (Fig. 8b).
Lastly, mean duration for individual Hamus
species is 16.8 + 9.4 years and the relative fre-
quency of species durations is skewed towards
shorter durations (Fig. 8a).
RELATIONSHIPS AND EVOLUTIONARY SCE-
NARIO.— Members of the ingroup are derived
from Hamus huxleyi by the acquisition of a
shell, pedal nerve cords, gonads, the movement of the gills into the mantle cavity, and the loss of
a differentiated foot, style sac, radula, jaws, aorta, statocysts, epipodium, buccal ganglia, optic ten-
tacles, and buccal mass. This grade in the evolution of Hamus (which is generally retained through
Hamus moolafisherorum) represents a sort of “average” mollusc, with emphasis upon the more
“primitive” forms. The basic body plan was arrived at by assuming that characters present in the
“most primitive” members of each (extant) class were also present in a common ancestor, and all
(extant) classes could be derived from it (see Huxley 1853). Thus, many early Hamus species were
intended to encompass the characters of the five molluscan classes then known. The discovery of
other taxa (such as the fossil Rostraconchia and the still extant Monoplacophora) has done surpris-
ingly little to induce serious rethinking of this original structure (see below).
The only apomorphy of the Lankester group is the appearance of both a posterior and anteri-
or aorta. The HAM of Nicholson and Lydekker (1889) traces its ancestry directly from Hamus
lankesteri, but shows little of the anatomical retrogression seen in Hamus swinnertoni and Hamus
eastoni. The lack of a radula and gonad in Hamus nicholsoni suggests a possible common ances-
tor, but the lack of tentacles and the cap-shaped shell also suggests a spurious relationship with
Hamus pelseneeri, which would not make an appearance for another 17 years. A peculiar autapo-
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 673
9.80 morphy characterizes Hamus nicholsoni — the
anterior mantle cavity with its single branchia
(Fig. 10b). This untorted mollusc with an ante-
rior mantle cavity would be a “hopeful mon-
ster” in anyone’s phylogeny, and although it
might have served as a novel preadaptation for
the Gastropoda, it never produced any descen-
ao creme dants. The HAM illustrated by Swinnerton
(1923) is directly descended from H. lankesteri
: j : : (“after Lankester”) and is the first instance of
ee eo eee ee ee eerie se PACMOMORPhOsISam the taxonmMamusHamuUs
b | swinnertoni (Fig. 9a) has lost numerous organs,
° including the radula, gonad, heart, pericardium,
°)| kidneys, and the entire nervous system.
Moreover, the shell of this species is conical
rather than cap-shaped, and the overt morphol-
ogy is distinctively gastropod-like, in spite of
its degenerate viscera. Hamus eastoni is the sis-
1 ee0e coo ee ter taxon of H. swinnertoni and is even more
°
w
oO)
Frequency (%)
10 #15 2 25
0.20 5 Taxon duration (yrs)
Taxa yr
F RA ieeh EG Dal. Leads, degenerate (Fig. 9b).
1860 1880 1900 1920 1940 1960 1980 2000 The common ancestor of Hamus pelse-
Year neeri and Hamus borrapottsorum is further
FIGURE 8. Macroevolutionary data for Hamus. a. extinc- | Characterized by the movement of the digestive
tion and origination rates, and species duration; b. species gland from the ventral to dorsal position,
diversity. gonads opening into the percardium, and the
loss of eyes. This basic body plan was arrived at by assuming that the characters are present in most
of the remaining taxa with the exception of members of the Stasek group. Something analogous to
“species selection” may help to explain why Pelseneer’s HAM had such a strong influence on
Anglo-American HAM speciation. Pelseneer was a Belgian and wrote mostly in French, but H.
pelseneeri appeared in a very influential textbook in English in 1906. Actually, this was a punctu-
ational event in a marginal habitat — Pelseneer (1897) published an earlier version in French.
Pelseneer’s HAM species also had an important dispersal episode. Pelseneer (1885) remarks that
he worked in Lankester’s laboratory during the winter of 1884-1885, thereby allowing us to doc-
ument not only direct ancestry, but perhaps the original dispersal event between England and the
continent as well. The HAM of Moore, Lalicker, and Fischer (1952) (Hamus moolafisherorum) 1s
diagnosed by ventral digestive glands, and the loss of the pericardium, gonads and pedal nerve
cords.
The remaining HAMs are divided into two distinct subclades, and it is this divergence between
1952 and 1958 that marks the beginning of the modern Hamus radiation (Fig. 6). Before this
branching point, the phylogeny of HAM was primarily comb-like (Fig. 6), the earlier HAMs form-
ing a grade of evolution. The synapomorphies that united these two subclades are the presence of
the osphradium and radula. Three synapomorphies diagnose the Morton group; all are typical gas-
tropod characters. They include the presence of a style and gastric shield, osphradium, and affer-
ent gill membrane. The five synapomorphies that diagnose the Stasek group are the reappearance
of gonoducts, osphradium positioned on dorsal surface of pallial cavity, multiple shell attachment
muscles, and the loss of cephalic tentacles and the digestive gland.
Like Hamus swinnertoni (Fig. 9a) the HAMs of Russell-Hunter (1968) (Fig. 9c) and Barnes,
674 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
Calow and Olive (1993) are undoubtedly products of paedomorphic processes that have beset these
species, and as with so many cases of degenerative evolution, we have at least two cases of con-
vergence amongst four taxa. While the known phylogeny of H. swinnertoni allows us to identify
its ancestry, we have no indisputable evidence for the determination of the relationships of the
remaining three species. However, mentorship distribution and stratigraphy provide insights into
their relationships (see discussion of the Morton group below).
The HAMs of Stasek (1972), Boss (1982), Raven and Johnson (1992) (Fig. 9d) and Salvini-
Plawen and Steiner (1996) are all flatworm-like. In addition, the solid shell is lost and replaced by
a spicule-studded integument in Hamus bossi, Hamus ravenjohnsonorum, and Hamus salvini-
steinerorum.
The appearance of the Morton Group marks a major evolutionary event within the genus (Fig.
6). Members of the Morton group have the most complicated anatomies of any of the Hamus
species, emphasize gastropod features, and mark a clear departure from previous HAM lineages.
This increasing convergence with the Gastropoda is clearly illustrated by the sequence a>b—c—d
in Fig. 10 (see also Haszprunar 1992).
Hamus mortoni is diagnosed by six autapomorphies (jaws, two digestive glands, epipodial ten-
tacles, salivary glands, esophageal glands, and spherical kidneys), all of which further emphasize
gastropod features. Many features present in the gastropod-like H. mortoni are absent in the
Morton and Yonge (1964) HAM. Hamus mortonyongorum has lost the osphradium, afferent gill
membrane and eyes. The nervous system is also condensed, and only a single ganglion remains.
Two American HAMs are basal members of the Morton group — Hamus barnesi (Barnes
1963) and Hamus meglitschi (Meglitsch 1967). Synapomorphies include the presence of the
osphradium on the efferent membrane of the gill and the loss of the hypobranchial gland. The most
unusual feature in this subclade is the knobs on the tentacles of H. meglitschi — evidently the par-
allel selection pressure toward gastropod structure in North America has turned it into somewhat
of a pulmonate gastropod!
Two more paedomorphs, Hamus russellhunteri and Hamus barcalolivorum, are treated here as
members of the Morton group. This placement is congruent with stratigraphic data and makes
sense considering the academic parentage of their creators.
Several non-Anglo-American HAMs often superficially resemble or are erroneously attributed
to insular Hamus species. For example, the South American HAM (Camacho 1966) resembles both
Hamus mortoni and Hamus mortonyongorum. However, this similarity is entirely due to conver-
gence. Camacho’s HAM was copied with virtually no changes from Portmann (1960: fig. 1470).
This, in turn, was a modification of a figure by Naef (1924), which, however, represented the
ancestral conchiferan, not the ancestral mollusc. Naef derived the molluscs from the annelids, and
Portmann reduced the gills from two pairs to one. There is also Dechaseaux’s (1952) HAM species,
which claims its ancestry as “after Lankester,” but is in fact identical to the paleo-paedomorph H.
swinnerton.
DISCUSSION
A major feature in Hammian evolution has been the expansion and enlargement of the poste-
rior mantle cavity and the migration of gills into it (Fig. 10). It is significant that the posterior man-
tle cavity, with its paired gills and associated structures and orifices, is not an ancestral, but a
derived condition. The canalization of HAM morphology also deserves special mention. In spite of
new discoveries in the field of malacology, such as the discovery of living Monoplacophora in the
1950s or the recognition of the Rostroconchia in the late 1970s, the general morphology of Hamus
species has remained little modified. Lineages that respond to changes in the environment tend to
be short-lived and quickly go extinct. This phenomenon is particularly well documented in the
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY
FiGureE 9. Paedomorphic and flatworm-like Hamus species. Paedomorphs include a. H. swinnertoni, b. H.
eastoni, c. H. russellhunteri, and the flatworm-like d. H. ravenjohnsonorum. Note virtual lack of mesodermal
structures (e.g., kidneys, heart, gonads). Redrawn from Swinnerton (1923), Easton (1960), Russell-Hunter
(1968), and Raven and Johnson (1992), respectively.
FiGurE 10. Generalized HAMs showing major trends in Hammian evolution over the last 150 years. Note
increasing complexity and number of organs in transition from a—d, and the teratological Hamus nicholsoni (b)
with its anterior mantle cavity, and yet untorted nervous system and alimentary tract. a. after Lankester (1883),
b. after Nicholson and Lydekker (1889), c. after Barnes (1963), and d. after Seed (1983) (from Hickman and
Lindberg 1985).
675
676 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
Morton lineage by Hamus mortonyongorum, Hamus barnesi, and Hamus yongethompsonorum. All
three of these species have serial pedal muscles, reflecting the discovery of the monoplacophoran
Neopilina. However, not one species that has originated since 1976 shows any segmentation what-
soever and only H. barnesi remains extant. HAM’s normal environment is a sort of pedagogical
refugium, in which degenerative reversions are quite common. Strong stabilizing selection may
also have helped to keep HAM on a maladaptive peak.
The Morton group is convergent with the Gastropoda in many aspects of its anatomy. One
wonders if some kind of mimicry is perhaps involved, though something more like lateral gene
transfer is perhaps a better way to put it. Neontological and paleontological HAMs are interspersed
throughout the early history of the group and have no intrinsic characters that distinguish them.
The evolution of HAM exhibits all the typical processes and developmental heterochronies
thought to encompass organic morphological evolution, and therefore both phenetic and cladistic
analyses have problems with paedomorphic taxa. Loss of organs and other features, particularly the
radula and gonads, is a major process in HAM evolution (there are no data on how HAMs that lack
gonads reproduce, but for modern species photocopying 1s a distinct possibility). Metamerism, or
the duplication of structures, occurs in the pedal musculature, gills, digestive glands, and is — sur-
prisingly —a derived rather than ancestral feature within the group. In many Hamus species,
organs appear in the juvenile condition, although the animal as a whole is represented as an adult.
Such paedomorphosis is seen in the various degrees of development of the nervous system. Loss
of the gonads is perhaps a case of progenesis, these being the last mesodermal structures formed,
but it 1s hard to see how such a change could not be a “lethal” mutation in terms of Darwinian fit-
ness. The most extreme form of heterochrony in Hamus species appears in Hamus swinnertoni,
Hamus eastoni, and Hamus russellhunteri. In these species only endoderm- and ectoderm-derived
structures are present. Mesoderm derivatives fail to develop, and the degenerative anatomy of these
species is readily apparent (Fig. 9).
THE INFLUENCE OF HAM ON MOLLUSCAN STUDIES
HAM’s fitness depends upon a symbiotic relationship with its pedagogical environment. It has
not aided evolutionary biologists in solving problems, but it has often had the opposite effect, by
requiring that theories be treated in the context of HAM. HAM has functioned as a Procrustean bed
in molluscan phylogenetics because its advocates have presupposed as an answer what ought to
have been the question. Namely, they have taken a phylogenetic hypothesis to be a fact, instead of
testing its merit relative to some alternative. With the advent and implementation of molecular
techniques over the last 10 years, many of the questions that HAM was inappropriately used to
address have diminished in their import. Here we examine two previous uses of HAM in evolu-
tionary debate — (1) determination of the sister taxa of the Mollusca, and (2) the anatomy and clas-
sification of Cambrian univalve molluscs. While the question of the sister taxon has moved from
the morphological to the molecular arena, HAM’s presence in the paleontological literature
remains problematic. And regardless of the state-of-our-knowledge, both examples provide valu-
able insights into the risks associated with hypothetical ancestors and their uncritical ‘evolution’ as
described above.
FLATWORMS OR ANNELIDS?— Prior to molecular data there were two major competing
hypotheses about molluscan origins: (1) molluscs are modified flatworms, (2) molluscs are modi-
fied annelids. Ghiselin (1988) and Winnepenninckx et al. (1994, 1995) provide some of the earli-
est analyses of small subunit ribosomal DNA (18S) to address this question. These studies, and oth-
ers, including Field et al. (1988), Lake (1990), Turbeville et al. (1991, 1992), have served as the
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 677
basis for many molluscan sister taxon comparisons, and they have consistently placed the Mollusca
among the lophotrochozoan taxa (molluscs, annelids, brachiopods, bryozoans, and phoronids).
However, the relative branching pattern for these taxa has yet to be resolved (Halanych et al. 1995).
For example, Zrzavy (1998), using a combined analysis of 18S data and morphology, suggested
that the sipunculids were the sister taxon of the molluscs. However, Boore and Staton (2002), using
partial mitochondrial gene order data, suggested the sipunculids are actually more closely related
to annelids rather than molluscs. Mallat and Winchell (2001), based on a 28S data set, suggested
that brachiopods and/or phoronids may be the molluscan sister group. In no case have the flat-
worms been supported as the sister taxa of the Mollusca, so why was the prior debate so persist-
ent? Was the morphological data so homoplastic and perplexing that previous workers were unable
to resolve relationships?
In making phylogenetic statements based on morphology (or molecules) one should proceed
as follows, compare the organisms, homologize as fully as possible, and treat the organisms shar-
ing the most synapomorphies as sister-groups. One might also want to use additional techniques,
but most people agree that we should at least do these things. If we do, it is clear that molluscs and
flatworms share many symplesiomorphies — e.g., spiral cleavage — which annelids also possess.
But molluscs and annelids also share many synapomorphies not present in flatworms — an anus,
a coelom, a particular larval type, just to mention a few. On this evidence, one has to conclude that
the molluscs share a more recent common ancestor with annelids than they do with flatworms. To
refute this line of reasoning, one must show that there are flatworm-mollusc synapomorphies not
shared with annelids. Historically, the only ones worth mentioning are some alleged features of the
pedal musculature, and hermaphroditism — the former evidently convergent and the latter demon-
strably so. Alternatively, one has to disestablish the mollusc-annelid synapomorphies.
These arguments were readily available decades before molecular data, but because the flat-
worm theory and its implications, including the unsegmented HAM, had been presupposed, the
annelid theory was widely dismissed. The arguments brought in to bolster an unsegmented HAM
and the flatworm theory have been a veritable museum of fallacious logic. We give eight examples.
(1) We have the logical fallacy of basing an argument upon negative evidence. Lack of evi-
dence that X occurred is taken as evidence that Y occurred. We are told that annelidan conditions
are not recapitulated, as in certain aspects of the development of Chiton (Hammersten and Runn-
str6m 1925). There are plenty of examples of non-recapitulation, for instance chicken-teeth. When
something is recapitulated, it is a fact to be explained, and nothing more.
(2) We have the formal fallacy of irrelevant conclusion. Russell-Hunter and Brown (1965)
assert that Neopilina’s structure does not fit Hyman’s definition of “metamerism.” The question is
not whether how somebody uses a word applies, but rather what has happened. The issue is not
whether molluscs “are” segmented, but what their ancestors were, and what their relationships are.
(3) We have rampant ad hoc hypothesizing. Many authors have been able to imagine reasons
why, say, Nautilus needs more gills (Hoffmann 1937), but they need additional ad hoc hypotheses
to explain away the multiple kidneys, and coelomoducts.
(4) We find a habit of treating the relational property “primitive” as if it were, like “spiny,” an
intrinsic one, to be read off without comparison. Korschelt and Heider (1900) argue, in so many
words, as follows: Cephalopods are advanced; Nautilus is a cephalopod; Nautilus has four gills;
therefore, having four gills is an advanced trait. It is curious that among the cephalopods Nautilus
is considered “primitive” — except in precisely those features in which it contradicts HAM and the
flatworm theory.
(5) We get a conflation of historical relationships and what we see in extant forms. Clark
(1980) claimed to have discovered the perfect example of metamerism in oligochaetous annelids.
678 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
Molluscs, he said, are different, and therefore are not derived from annelids. This amounts to belief
in a physiological essence. Segmentation, like all sorts of other things, has evolved. There is no rea-
son to believe that the physiologically ideal state represents the ancestral one, and even if it did, it
would not show a lack of descent.
(6) We find the possibility of an event having occurred treated as if it had in fact occurred.
Clark, again, shows that metamerism has evolved independently among tapeworms, chordates, and
the annelid stock. If not three times, why not any number at all? The opposite conclusion accords
with the principles without which it is impossible to infer relationships at all.
(7) We have circular reasoning, or begging the question, in which attempts to support the flat-
worm model presuppose that the flatworm theory is true. Particularly in the works of Stasek (1972),
we find it asserted that molluscs are “pseudometamerous.” Why not “‘vestigially metamerous?”
(8) And, finally, we have an abuse of scenarios. There is some healthy controversy as to what
role scenarios ought to play in phylogenetic research. But if one is to use them, one ought to do so
logically. A logically legitimate role that they can play is in testing hypotheses. If a theory implies
the existence of hypothetical organisms that would not have been viable, that theory is false. But it
is the logical fallacy of denying the antecedent in a conditional statement, to “confirm” hypotheses
by showing that they do not contradict particular theses. Thus, both Stasek (1972) and Salvini-
Plawen (1980) provide us with a long account of what happened physiologically as flatworms were
transformed into molluscs — never considering the obvious fact that the annelid theory can accom-
plish the same end, without the long list of imaginary intermediates and numerous convergences
that the flatworm theory and HAM demand.
Of course, there is no real evidence that HAM ever existed. Indeed, the non-conchiferan mol-
luscs (aplacophorans and chitons) have been treated as “non-molluscs” by some authors because
they lack the “defining characters” found in HAM (e.g., Odhner 1919), and the lack of an HAM
stage in the ontogeny of any extant mollusc was first noted by Verrill (1896).
Today, the “flatworm versus annelid” controversy is being argued in somewhat different terms.
Some authors argue that body ‘segmentation,’ upon which the supposed relationship of the mol-
luscs, annelids, and arthropods was largely based, is actually convergent. In parallel, our under-
standing of fine structure, development, and “segmentation” of both molluscs and annelids has also
markedly increased with the application of modern imaging and developmental techniques as well
as detailed anatomical studies of basal molluscan groups (e.g., Salvini-Plawen and Bartolomaeus
1995; Haszprunar and Schaefer 1997; Wanninger and Haszprunar 2002). Others, however, main-
tain very much the opposite thesis, namely that segmentation has been secondarily reduced not
only in molluscs but in many other groups and may even go back as far as the bilaterian common
ancestor (Balavoine and Adoutte 2003). Part of the evidence comes from molecular trees, which
show that some animals with no obvious trace of segmentation are closely related to those with it.
In some of these, such as echiurans, there are morphological traces of segmentation as well
(Hessling 2002). The mechanisms that control the morphogenesis of segments in all animals have
been widely homologized as well. There may, however, have been a considerable amount of par-
allelism involved, so that the seriality was not as widely expressed as it is, for example, in modern
arthropods and annelids. At the very least, given that molluscs and annelids are more closely relat-
ed to each other than to arthropods, the kind of segmentation that has been hypothesized to have
been present in the common ancestor of annelids and arthropods must either be convergent or have
been secondarily reduced in Lophotrochozoa. Two theoretical points are worth mentioning in this
connection. In the first place, the notion that evolution always proceeds from simple to complex
was very common in the nineteenth century, and it is about time that it no longer be presupposed
in efforts at phylogenetic reconstruction. Second, there are good theoretical reasons for thinking
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 679
that it is easier for parts to be lost than gained. Our views on such matters may profoundly affect
our conclusions, but they are rarely made explicit. Although the ultimate outcome of the “segment-
ed common ancestor” question for the Mollusca remains elusive, we need to proceed with care and
rigor in testing alternative hypotheses and not rely on imaginary creatures to parse and test data as
in the past.
USE AND ABUSE OF FossiL HAMsS.— Paleontologists often try to stuff HAM into shells as if
they were dishing out escargot. Following a tradition that goes back at least as far as Knight (1952),
Peel (1974, 1991) and others have loaded large bellerophont shells onto tiny HAMs. The alterna-
tive that at least some bellerophonts were slugs was well argued by McLean (1984). Others, such
as Knight (1952), Pojeta and Runnegar (1976), and Peel (1990, 1991), tested alternative morpholo-
gies (i.e., torted or non-torted) for extinct Paleozoic molluscs, based on the fit of fossil conchs,
resplendent with holes, tubes, slits, and trails, on HAM’s anatomical body plan. Here we have real
data being “tested” against a hypothetical anatomy to determine whether the hypothesis shall be
accepted or rejected on the grounds of which reconstruction “looks comfortable.” Paleontologists
may claim to base their inferences on the fossil record and the structure of extant organisms. But
in the case of molluscs, their procedure all too often has been to use an imaginary organism —
HAM — as a Procrustean bed, to which the soft parts of fossils are fitted by adding features here,
lopping off others there, and stretching the data wherever they fall short of the desired effect.
Because of the use of HAM to distinguish untorted molluscs from torted gastropods, it has sur-
reptitiously become a linchpin in some torsion scenarios. Ghiselin (1966) suggested a possible sce-
nario for the non-saltatory origins of gastropod torsion. Batten, Rollins and Gould (1967) countered
with claims based upon unpublished results. These results, subsequently published by Rollins and
Batten (1968), employed HAM to justify the non-torted nature of the mollusc used to argue against
Ghiselin’s scenario. And, although Harper and Rollins (1982) ultimately retracted their claims, the
damage had already been done, and this episode has been used to discredit functional thinking.
Sanitation issues are often evoked in torsion scenarios because many of the holes, tubes, slits,
and trails of the fossil conchs are frequently linked with hypothetical water circulation patterns in
the HAM anatomy so that waste products will be removed from the mantle cavity. It 1s interesting
to note that Pelseneer (1894), who first discussed the sanitation problems that molluscs face with
the anus and gills in close proximity, also was the first to pen a HAM with the gills located within
the mantle cavity! Sanitation problems for Hamus species were intensified in 1952 when a new
character state appeared in the common ancestor of Hamus moolafisherorum and _ the
Morton/Easton groups (Fig. 5). This synapomorphy was the placement of the anus above the gills
rather than below them (i.e., downstream) (Appendix, character 20); so much for intelligent design.
Although potential outgroups such as chitons and monoplacophorans have the anus and gills sep-
arate and the anus located below the gills, Hamus evolution has not converged with these function-
ing anatomies and, instead, has been directed towards an ever increasingly maladaptive state.
Because of its more problematic nature, these maladaptive characters in the HAM phylogeny pro-
vide a much larger arena for speculation and interpretation and are vastly more interesting than the
character states found in real organisms.
SUMMARY
The difficulties associated with reconstructing relationships, hypothetical taxonomic units,
fossil anatomies, and incorporating fossil taxa into evolutionary scenarios by no means implies that
we should give up. Rather, we should try alternatives, especially alternatives to HAM. Many of our
colleagues have argued that HAM should be allowed to exist as a sort of pedagogical fairy tale, jus-
tified as a means of teaching molluscan anatomy. They acknowledge that no such creature ever
680 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
existed and that any evolutionary scenario deriving molluscan taxa from it represents misinforma-
tion. Such arguments remind us of ones given in favor of paraphyletic taxa. Unfortunately, neither
the imaginary animals nor the misleading groups come clearly labeled with warnings about the
harm that they might do if mistaken for real organisms or monophyletic units.
It is our conclusion that HAM is a pest being preserved in a textbook refugium, and science
needs a better basis for determining the structure, relationships, and classification of organisms
than an expedient of didactics and pedagogy. The sooner all Hamus species become extinct the bet-
ter.
ACKNOWLEDGEMENTS
This paper was begun in collaboration with the late D.P. Abbott forty years ago. It has been
presented to various audiences since 1965 and has benefited from many thoughtful discussions
with our colleagues through the years. We thank T. Gosliner, J. Griesemer, G. Haszprunar, C.
Hickman, and the anonymous reviewers for comments on the manuscript, and L. McConnaughey,
M. Taylor, and P. Spowart for preparing figures 2, 7 and 8, respectively.
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Appendix
DATA MATRIX FOR HAMUS SPECIES
Character Number
C)ONOVO KO) COO)OKO) ababscaababalabalal BAAVAZQZ2ZIQA 33332333
Taxon 123456789 0123456789 0123456789 0123456
H. huxleyi 202000000 00000000?0 0000700000 0?00000
H. lankesteri 010010000 0111000002 0021000010 0?11100
H. nicholsoni 11ITOOOTIO OLLIOOO002 003 OF ODOM OS ameROlO
H. pelseneeri TTLOLOLOO -OLTO OOO? 2 OOD1 1 OM TA Oso 2aeOlO
H. swinnertoni 111010100 0111000012 02°?70??0110 0O?14011
H. borrapottsorum LATO OAOO! OA LAO OO OAD OO 2AM OOM OROK AZ a eROlO
H. moolafisherorum 111010100 0111000002 1110770110 0224000
H. mortoni LATOLOLOO LOLOLIOO[ZL, LOU TA ODA OZ Ako
H. eastoni 1TOOLOLTOO O1LTOOO0122 1270? VOM MO OyemrarORtal
H. barnesi PATIL O A TAT OCOLOIL O21 VOD NOM si EA a Ole
H. mortonyongorum PATTI LOLOO OMT OOO OOM WOMAN 160 eRe OV 24 tO
H. hickmani TITOLOLOO LATOOOLOT2 LOD A OO ail ae ORRA RRO
H. meglitschi LTILOLVOLOO LLLITOLOL2 HOD 1a OO Aloe ime Amr OlO
H. russellhunteri TALOUOLOO WTLVOOOM?22 12°20? 201M ORs aOnml
H. staseki LAITTALTOOO OLLOOOOL?2 OOLLO CTA O. OFA anon
H. yongethompsonorum TTTITOLOO LILVOLOOOLT LOT PO dae eZ AA OO
H. barthbrosorum 1LIOLOLOO OLLOOOLOL2 102110021, Shoe aoe
H. bossi P2211 O0L00 OLLOOOOI22 1O2AO OM MO REZ AZ akOlo
H. seedi TALOLOLOD LLOOOOLO2AL DOMLOODLIAL MoOZA4aKolo
H. barcalolivorum LTLOLOLOO OLLOOOVVOLE HODLLOODTOMOR 2S MOO
H. ravenjohnsonorum 2222710100 0110000122 12270? ?1 1109 02240nm
H. salvinisteinerorum 227110100 OLLO000L?22 LOLIT0OLI TO i2240ne
“Cambrian” taxa
H. pojetarunnegarorum species complex:
Taxon a LITOLOLTO OLTIOOOL?2 O21T0? 20110 OR2 Aone
Taxon b LILOLOL2ZO OLTIOOOL?22 O21TO? 2010 Oe 2A One:
Taxon c 1IOOLOI1IO OLLIOO0LT?22 O20 2? O1LO OPA Onm
Taxon d LLOOLOI20 OLLTLIOOOL?2 0210720110 ,0R2a0nar
Taxon e LLOOLOLLO OLLLOOOL?22 1220? POO MORZAOne
Taxon f TLOOLOII20. OLLLOOOT22 2250? eos OR AAO ne
LINDBERG AND GHISELIN: MOLLUSCAN ARCHETYPES AND PHYLOGENY 685
CHARACTER ANALYSIS
Integument
1. Shell enveloped by mantle. The plesiomorphic state is equivocal as the outgroup lacks a shell. States —
shell enveloped by mantle = 0, shell not enveloped by mantle = 1.
2. Exoskeleton. Shell absent is the plesiomorphic state. States — shell absent = 0, shell present = 1, integument
with spicules = 2.
3. Shell with opening. The presence of an opening in the shell is equivocal. States — shell opening present =
0, shell without openings = I.
4. Shell attachment muscles. The absence of multiple muscles is plesiomorphic. States — multiple muscles
absent = 0, multiple muscles present = 1.
Foot
. Foot divided into propodium, mesopodium and metapodium = 0, foot not differentiated = 1.
Gills
6. Number of gills. A single pair of gills is plesiomorphic. States — single pair of gills = 0, multiple gills = 1.
7. Gills enclosed in a mantle cavity (internal) or extending beyond body (external). External gills are ple-
siomorphic. States — external gills = 0, internal gills = 1.
8. Gill position on body. Gills situated on the posterior portion of the body are plesiomorphic. States — pos-
terior placement of gills = 0, anterior placement of gills = 1, lateral placement of gills = 2.
9. Efferent membrane. The absence of an efferent membrane from the gill to the roof of the mantle cavity is
plesiomorphic. States — gill without efferent membrane = 0, gill with efferent membrane = 1.
10. Afferent membrane. The absence of an afferent membrane from the gill to the floor of the mantle cavity
is plesiomorphic. States — gill without afferent membrane = 0, gill with afferent membrane = 1.
On
Digestive System |
11. Jaws. The presence of jaws in the buccal cavity is plesiomorphic. States — jaws present = 0, jaws absent
= il
12. Buccal mass. The presence of a buccal mass is plesiomorphic. States — buccal mass present = 0, buccal
mass absent = 1.
13. Radula present in oral cavity. The presence of a radula is plesiomorphic. States — radula present = 0, radu-
la absent = 1. ;
14. Salivary glands. The absence of salivary glands is plesiomorphic. States — salivary glands absent = 0, sali-
vary glands present = 1.
15. Esophageal glands. The absence of esophageal glands is plesiomorphic. States — esophageal glands
absent = 0, esophageal glands present = 1.
16. Configuration of intestinal tract. A straight, non-looped intestinal tract is plesiomorphic. States — intes-
tinal tract straight = O, intestinal tract looped = 1.
17. Digestive gland. The presence of a digestive gland is plesiomorphic. States — digestive gland present =
0, digestive gland absent = 1.
18. Position of digestive gland relative to the stomach. A ventral digestive gland is plesiomorphic. States —
digestive gland ventral = 0, digestive gland dorsal = 1, both dorsal and ventral digestive glands = 2.
19. Style. The presence of a style sac without a style is plesiomorphic. States — sac without style present =
0, style present in stomach = 1, both sac and style absent=2.
20. Anus position. An anus opening below the gill is primitive. States — anus positioned below the gill = 0,
anus positioned above the gill = 1.
686 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, No. 27
Coleomic structures
21. Heart and pericardium. The presence of a heart is plesiomorphic. States — heart present = 0, heart and
pericardium present = 1, heart absent = 2.
22. Aorta. The presence of an anterior aortic branch is plesiomorphic. States — anterior aorta only = 0, aorta
absent = 1, anterior and posterior aorta = 2, posterior aorta only = 3.
23. Gonads. The absence of gonads is plesiomorphic. States — gonads absent = 0, gonads present = 1.
24. Gonoducts. Separate gonoducts are plesiomorphic. States — separate gonoducts = 0, gonads opening into
pericardium = 1.
25. Kidney morphology. Tubular kidneys are plesiomorphic. States — kidneys tubular = 0, kidneys spherical
= Il,
Sensory structures
26. Cephalic tentacles. The presence of cephalic tentacles is plesiomorphic. States — cephalic tentacles pres-
ent = 0, cephalic tentacles absence = 1.
27. Statocysts. The presence of statocysts is plesiomorphic. States — statocysts present = 0, statocysts absent
= ll.
28. Epipodium and tentacles. The presence of an epipodium without tentacles is plesiomorphic. States —
epipodium present = 0, epipodium absent = 1, epipodial tentacles present = 2.
29. Hypobranchial gland. The absence of a hypobranchial gland is plesiomorphic. States — hypobranchial
gland absent = 0, hypobranchial gland present = 1.
30. Osphradium. The absence of an osphradium is plesiomorphic. States — osphradium absent = 0, osphra-
dium present = |.
31. Osphradium position. The position of an osphradium on dorsal surface of foot is plesiomorphic. States —
osphradium on dorsal surface of foot = 0, osphradium present on efferent membrane = 1, osphradium pre-
seent on dorsal surface of pallial cavity = 2.
32. Eyes. The presence of stalked eyes is plesiomorphic. States — stalked eyes = 0, non-stalked eyes = 1, eyes
absent = 2.
Nervous system
33. Ganglia. The presence of four pairs of ganglia (cerebral, pedal, parietal, buccal) is plesiomorphic. States
— all four pairs of ganglia present = 0, three pairs present (cerebral, pedal, parietal) = 1, cerebral only =
2, cerebral + pedal = 3, ganglia absent = 4.
34. Pedal nerve. The absence of pedal nerve cords is plesiomorphic. States — pedal nerve absent cords = 0,
pedal nerve cords present = |.
35. Visceral nerve loop. The presence of a visceral nerve loop is plesiomorphic. States — visceral nerve loop
present = 0, visceral nerve loop absent = 1.
36. Nerve ring. The presence of a nerve ring around the pharynx is plesiomorphic. States — nerve ring around
the pharynx = 0, absence of nerve ring around the pharynx = 1.
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
687
(oy)
ACKNOWLEDGMENTS
The Editors of the Academy’s Scientific Publications wish to acknowledge, with
thanks, the efforts of the many reviewers who have given unstintingly of their time
and knowledge to review papers submitted for publication in the Academy’s
Proceedings volume 54 for year 2003. The following persons provided critical
expertise: Michele L. Aldrich (Cornell University), Frank Almeda (California
Academy of Sciences), K. Christopher Beard (Carnegie Museum of Natural
History), David Behrens (Schafer Laboratories), Hans Bertsch, Daniel M. Cohen
(Bodega Bay, California), Thomas F. Daniel (California Academy of Sciences),
Indraneil Das (University of Malaysia), Terry Erwin (National Museum of Natural
History, Smithsonian Institution), Lawrence J. Flynn (Peabody Museum of
Archaeology and Ethnology, Harvard University), Darrel Frost (American Museum
of Natural History), Terrence M. Gosliner (California Academy of Sciences),
Gustavo Hormiga (George Washington University), Tomio Iwamoto (California
Academy of Sciences), David H. Kavanaugh (California Academy of Sciences), J.
Patrick Kociolek (California Academy of Sciences), John E. McCosker (California
Academy of Sciences), Nigel R. Merrett (UK), Richard J. Mooi (California
Academy of Sciences), Timothy Pearce (Carnegie Museum of Natural History),
Richard L. Reeder (University of Tulsa), William Rudman (Australian Museum),
William Shear (Hampden-Sydney College), Carol Tang (California Academy of
Sciences), Fred G. Thompson (Florida Museum of Natural History, University of
Florida), Angel Valdes (Los Angeles County Museum), Jens V. Vindum (California
Academy of Sciences), Jeff Wilkinson (California Academy of Sciences), Gary
C.Williams (California Academy of Sciences), Guinevere O.U. Wogan (California
Academy of Sciences), Ellis L. Yochelson (USGS and National Museum of Natural
History, Smithsonian Institution), George R. Zug (National Museum of Natural
History, Smithsonian Institution).
Alan E. Leviton, Ph.D., Editor
Gary Williams, Ph.D., Associate Editor
14 November 2003
ca ped pet pet pel pel fel pel pel pel pel pel pepe fel pel ped fed ped ped pe ped ped fed ped fed fe ed fed ed ed fed fed ped fed fed fe fe fed ped pe
fl
INDEX TO VOLUME 54
A
Acanthaceae 371, 379-380
Acanthemblemaria castroi 159
Acanthophora 307-309, 314, 318; A. spicifera
307-308
Actinicyclus [sic] 50
Actinocyclus 43-44, 58, 205; A. curvatulus 43, 58; A.
divisus 43; A. ehrenbergii 43; A. ochotensis 43, 58;
A. octonarius 43, 58
Actinoptychus 44, 58; A. senarius 44, 58; A. splendens
44, 58; A. undulatus 44; A. vulgaris 44
Adoneis pacifica 40, 44, 50
Aeolidina 161
Afghanistan 151
Africa: See countries
Afroedura hawequensis 236
Agamidae 1, 15, 16
Agelena ignota 566, 568
Ahaetulla 407
Ailuropoda 212-214, 220-221
Ailuropoda-Stegodon fauna 212
Akysis 85
Alaska 27-29, 31, 33, 35, 37-43, 45, 47, 49, 51, 53,
S550 7.59, Ol 63; 262. 310, 329; 4375 Alaska
Peninsula 27—29, 40; Kodiak Island 310
Alloiodoris 204
Amaurobiidae 499, 576, 577, 578
Ambanus 575-576
Amolops 21
Amphiliidae 81-83, 86-87, 89, 92, 98, 107-110,
113-114
Amphiliinae 83, 87-88, 108-110
Amphiesma 407
Amphilius 81-83, 87, 89, 97, 101, 103, 105-106, 108,
114; A. angustifrons 81-82, 97, 105; A. atesuensis
87; A. baudoni 87; A. brevis 87; A. cryptobullatus
87; A. kivuensis 87; A. lentiginosus 87; A. lon-
girostris 87; A. maesii 87; A. notatus 81-83, 106;
A. opisthophthalmus 87; A. pictus 87; A. ura-
noscopus 87; A. zairensis 87
Amphora bigibba var. capitata 246, 253
Anapidae 359
Anapis 359
Anapisona 359
Anapistula 356, 360; A. caecula 356, 360
Andersonia 83, 87-88, 94, 109, 117; A. leptura 83,
87, 94, 109
Andaman and Nicobar Islands: See Indian Ocean
Angola 86, 95, 113
Anguis platura 437
Anisodoris 71, 74
Anura 16, 17, 26, 141, 152-153
Apodemus 213
Arachnoidiscus 44
Araneae 133, 140, 356, 360, 499, 576, 577, 578, 579
Araneoidea 356, 360
Aristochroa 238-239, 241-244; A. abrupta sp. nov.
238-244; A. dequinensis 244; A. dimorpha
239-240; A. gratiosa 239-240; A. kangdingensis
239-240; A. militaris 239-240, 244; A. sciakyi
239-240
Asiacoelotes 502, 507, 561, 564, 576
Aspis intestinalis 423
Astarte 27-31, 39, 42
Asteronotus 205
Atlantic Ocean 381, 388; See also Cape Verde
Archipelago; See also Madeira
Aturia ornata 433
Auchenoglanidinae 82, 89, 109
Aulacodiscus 44
Auliscus 38, 44; A. grunowii 44
Australia 189-191, 204, 207-208, 231, 262, 280,
283-284, 286-288, 296-300, 303, 321, 323,
328-331, 356, 361, 366, 393-394, 403, 405406,
428, 430, 432, 434, 436-437, 450, 496; Kangaroo
Island 393-394; New South Wales 186, 190-193,
Dal 287% 296; 9300553035 23 6k 3660503 69:
Queensland 206, 323; South Australia 393-394,
405; Victoria 303, 330, 361, 374, 379; Western
Australia 190-191, 208, 284, 286, 296, 303, 323,
331; See also Tasmania
Axis 162, 221, 260, 363, 367
Azemiopinae 407, 438
Azemiops 407, 413, 419, 438-439, 450, 452; A. feae
413, 419, 438-439, 450
Azpeitia 38, 44; A. nodulifera 44; A tabularis 44
Azurina eupalama 159
B
Bacteriasrtum [sic] 44; B. varians 44 |
Bacteriastrum 38, 52; B. varians 38, 52
Bacterosira 34, 37-38, 40, 44, 54, 56; B. fragilis 34,
37-38, 44, 54, 56
Bagarius 85, 110, 113
Bagridae 82-83, 86, 88, 108-109, 113-114
Bagrus 109
689
690
Bangladesh 1, 11, 13, 425, 435, 440, 444, 446
Baptodoris 204, 205
Belonoglanis 83, 87
Bay of Bengal: See under Indian Ocean
Bering Sea 27, 38, 41-42
Bering Strait 27-28, 39, 40-42
Berthella 205
Bhutan 507, 521, 522, 548-549, 555, 576
Bifidocoelotes 499-500, 502-503, 507, 561, 564,
576, 581, 645-646; B. bifidus 502, 581: B. primus
502-503, 581, 645
Biomphalaria glabrata 229, 230
Bitia 407
Boiga 407
Borneo 294, 296, 422, 435, 438, 451
Bos 212, 213-214, 217
Bovidae 219
Brazilios, 715 73, 79
Bubalus 213, 217, 220
Bufo 11, 16, 141-153; B. andrewsi 150; B. asper 145,
150; B. atukoralei 145; B. bankorensis 145; B.
beddomi 145; B. biporcatus 145; B. bufo 145, 150,
153; B. burmanus 150; B. calamita 145; B. clav-
iger 145; B. crocus sp. nov. 142—146, 149-150; B.
cyphosus 145; B. dhufarensis 145, 151; B. diver-
gens 145; B. dodsoni 145; B. gargarizans 145,
151; B. himalayanus 145, 151; B. japonicus 145;
B. juxtasper 145; B. kisoloensis 145; B. kotagamai
145; B. latastei 145; B. leutkeni 145; B. luristani-
ca 145; B. macrotis 144, 150-151; B. melanostic-
tus 141-142, 145-152; B. melanosticus [sic] 145,
150; B. microtympanum 145, 151; B. minshanicus
145; B. noellerti 145; B. olivaceus 145; B. orien-
talis 145, 151; B. parietalis 145; B. parvus 145,
150-151; B. pentoni 151; B. periglenes 145; B.
peripatetes 145; B. philippinicus 145; B. quadri-
porcatus 145; B. raddei 145, 151; B. scaber 145,
151-152; B. silentvalleyensis 145; B. stejnegeri
145, 152; B. stomaticus 145, 148-149, 151; B. stu-
arti 16, 141-142, 144-146, 148-151; B. surdus
145, 151; B. tibetanus 145; B. viridis 145, 152: B.
wrighti 145
Bufonidae 16, 141, 149, 152-153
Bungarus 407, 409-411, 413-415, 421-423, 452,
453; B. annularis 421; B. bungaroides 411, 415,
421; B. fasciatus 411, 415, 421; B. flaviceps 409,
415, 422; B. magnimaculatus 415, 422-423: B.
caeruleus magnimaculatus 422: B. multicinctus
409, 423; B. multicinctus wanghaotingi 423; B.
wanghaotingi 407, 409, 413, 415, 423
Burma 422-423, 438, 443, 451, 463, 465, 494, 496,
497; See also Myanmar
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
C
Cadlina 205
Caelorinchus 281-282, 288
California 68
Calliophis 414-415, 423-424, 427, 452-453; C.
bivirgatus 414; C. gracilis 423; C. macclellandi
427; C. maculiceps 415, 424
Callophis maculiceps 424
Caloneis 245-246, 253: C. disticha 245; C. liber var.
incerta 246, 253
Calotes 1-11, 13-15; C. chincollium sp. nov. 2, 4,
6-13: C. emma 1\—2, 10-13; C. emma alticristatus
10-11; C. jerdoni 1-2, 10-11, 14; C. kinabaluen-
sis 1; C. kingdonwardi \-3, 11, 14; C. kingdon-
wardi bapoensis 14; C. maria 2, 40; C. mystaceus
1-2, 10, 12, 14; C. nigriplicatus 1; C. versicolor
1—2, 10, 11
Calycidoris 204-205
Cambodia 422, 424, 425, 442, 443, 445, 447
Cameroun 94, 95, 113
Campylodiscus cordatus 246, 253
Canis 217, 219-220, 223
Cape Verde Archipelago 381, 387
Cantoria 407
Caprinae 213
Carabidae 238, 244
Carminodoris 169-171, 173, 175-177, 179, 181, 183,
185, 187, 189; 191, 193, 195, 197, 19952012207;
C. armata 202, 206; C. bifurcata 169, 185, 189; C.
grandiflora 169-170, 175, 177-178, 194, 205; C.
mauritiana 169-170, 176-178, 194, 205: C. nodu-
losa 169, 175, 185, 189, 204-205
Cenozoic 27—28, 31, 40-42
Cerberus 407
Cervidae 213, 219-220
Cervus 212-213, 216-217, 220-221, 223
C. cf. unicolor 217
Chad Basin 110
Chaetoceros 30, 38, 44-45, 54: C. cinctus 44; C.
diadema 44; C. furcellatus 44; C. incurvus 44; C.
septentrionales 44; C. subsecundus 44
Cheilinus 361, 369
C. arenatus 361
C. coccineus 369
C. rhodochrous 369
Chiloglanis 85, 97, 104, 113
C. reticulatus 97, 104
Chilotherium yunnanensis 221
China 14, 24-26, 31, 133-134, 136, 138-139,
150-151, 209-211, 213, 215, 2ig Zio eae
223-224, 238-239, 241-244, 284, 286-287,
INDEX TO VOLUME 54
290-291, 295, 356-360, 409, 419, 422-423, 425,
427, 431-432, 434, 436, 438-446, 448-451,
501-509, 517-549, 555-579, 638-639; Anhui
Province 151, 449, 521-524, 538, 548, 550,
573-575; Fuji 568, 576; Fujian 449, 560; Gansu
449, 537-539, 544, 550, 575-576; Guangdong
449, 576; Guangxi 439, 449, 546-547, 556-557,
Se» Guizhou 0521563," aWile572;, 574—575,
638-640; Hong Kong 503, 576; Hubei 449, 501,
523, 525-526, 538, 540, 554-555, 563-564, 566,
571, 575-578, 606; Hunan 449, 501, 504-506,
518-519, 528, 536, 538-539, 549, 551-552, 559,
561-562, 565-566, 571-573, 576, 578, 636-637;
Jiangsu 449, 560, 566, 575-576; Jiangxi 449, 539,
573; Jilin 449, 567, 575; Ningxia Hui Autonomous
Region 151; Shaanxi 439, 522, 527, 533, 538, 555,
575; Shanxi 555-556, 575; Sichuan 425, 449, 505,
533, 550-553, 562-563, 565-566, 572, 575-576,
578: Szechwan 441, 443: Taiwan 25-26, 409, 429,
432, 434, 436, 440-442, 444, 448-451, 501-503,
537, 557-559, 576, 578; Tangzigou 209-210, 217,
224; Tengchong 221; Wanrengang 209-210,
215-217, 219, 224; Xizang Autonomous Region
(Tibet) 441, 442, 443, 446, 451, 577; Yangyi 210,
221, 224: Yunnan Province: 10, 14, 24, 133-134,
136, 138-139, 150, 209-211, 213, 215, 217, 219,
221, 223-224, 238-239, 241-244, 356-357, 360,
A409, 423, 425, 439, 441-443, 449, 496, 517-521,
524, 526-532, 535-536, 540-543, 545-549, 575;
Dianjiangtai 210, 221; Gaoligongshan 209-210,
223, 238, 242, 356-359; Gaoligong Mountains
133, 139, 209, 357, 360; Hengduan Range 209;
Huoxinshan 210, 217, 220, 224; Jiangdongshan
210, 221; Laohudong 209-210, 219-220, 222,
224; Longwangtang 210, 223-224; Nanfeng
209-210, 212-215, 224; Zhejiang 439, 449,
506-507, 520, 525, 534, 536, 538, 547-548,
553-554, 559-560, 575-577
Chioraera leonina 310
Chirixalus 11, 16-26; C. cherrapunjiae 22; C. doriae
17, 19-25; C. dudhwaensis 24, 26; C. eiffingeri
23, 25-26; C. hansenae 24; C. idiootocus 23; C.
laevis 22, 24-25; C. nongkhorensis 17, 19-21,
24-25; C. palpebralis 22-25; C. punctatus sp.
noy. 18-24; C. romeri 22; C. shyamrupus 22, 24;
C. simus 24; C. vittatus 17, 19-20, 23-25
Chiton 677, 683
Chromodoris 167, 204—205
Cladogramma dubium 44, 62
Clarias 85
Clariidae 109
Claroteidae 82, 86, 108-109
691
Claroteinae 82, 89
Clinus 156, 159; C. macrocephalus 159
Cocconeis 44; C. antiqua 44; C. californica 44; C.
costata 44; C. placentulla 44; C. pribiloeformis
44; C. scutellum 44; C. singularis 246, 253: C. vit-
rea 44
Coelocephalus 281, 283; C. acipenserinus 281, 283
Coelorhynchus 281-283; C. gladius 281
Coelotes 500, 502-510, 518-528, 531-541, 543-544,
546-569, 572-579, 594; C. acidentatus 518; C.
adligansus 518; C. altissimus 519; C. amygdali-
formis 520; C. arcuatus 520; C. argenteus 521; C.
aspinatus 521; C. atropos 575; C. charitonovi 575;
C. baccatus 540; C. baronii 521; C. bicaudatus
566-567; C. bifida 502; C. bituberculatus 522; C.
brunneus 523; C. calcariformis 523; C. carinatus
524; C. chaigiaoensis 525; C. cheni 525; C. coe-
datus 574; C. corasides 568-569; C. corasoides
566, 568; C. coreanus 526, 594; C. edentulus 558;
C. erraticus 566, 567; C. exitialis 575; C.
huangsangensis 503-504; C. huizhouensis 509; C.
huizhuneesis 509; C. icohamatoides 562; C. ico-
hamatus 562; C. ignotus 568; C. impletus 561; C.
kulianganus 560; C. lama 509, 533; C. laoyingen-
sis 564-566; C. latus 557; C. lichuanensis 563; C.
lutulentus 538; C. lyratus 572; C. magniceps 539;
C. michikoae 569; C. modestus 568, 577; C. mol-
lendorffi 560; C. molluscus 539; C. nanyuensis
539; C. neixiangensis 540; C. ornatus 541; C.
penicilatus 510, 543; C. penicillatus 543; C. per-
vicax 543; C. platnicki 557; C. pseudoluniformis
558-559; C. pseudoterrestris 575; C. gingzangen-
sis 546; C. quadratus 546; C. rufuloides 500, 509,
547-548; C. rufulus 547; C. saxatilis 510, 525; C.
schenkeli 548; C. senkakuensis 560; C. shuang-
paiensis 500, 510, 528; C. singulatus 549; C. sin-
ualis 538; C. stemmleri 549; C. streptus 550; C.
striolatus 550; C. strophadatus 550; C. subtitanus
551; C. syzygiatus 551; C. tautispinus 573; C. ter-
ebratus 551; C. trifasciatus 552; C. tryblionatus
553; C. uncinatus 553; C. urumensis 565; C. var-
iegatus 573; C. wenzhouensis 554; C. wudangen-
sis 554; C. wuermlii 555; C. yadongensis 556; C.
yoshikoae 503, 504; C. yosiianus 556; C. zonatus
564-565
Coleoptera 238, 244, 281
Colubridae 407
Coluber: C. gramineus 444; C. laticaudatus 425; C.
naja 424; C. russelii 439
Congo Basin 81-84, 86-87, 95-96, 99, 104, 110-112,
118; Stanley Pool 90-94, 97, 100, 102, 111, 113,
117, 119-122, 124-127, 129-130; Ubangui River
692
84, 88, 90-92, 97, 102, 104-107, 111, 118-119,
128, 130-132
Cook Islands: See South Pacific
Coral Sea: See Indonesia
Coras 568, 574, 578; C. lamellosus 574; C. luctuosus
568, 578
Coronilla 499-500, 503-506, 557, 576, 582-585,
646; C. gemata 499, 503-505, 582; C. huangsan-
gensis 505; C. libo sp. nov. 499, 503-505, 563,
574, 583; C. mangshan 503-505; C.. sigillata
503-506, 584; C. subsigillata sp. nov. 499,
503-504, 506, 585; C. yanling 499, 503-504
Coryphaenoides 279-280, 282, 293-294, 300-301;
C. hyostomus 293-294
Coscinodiscus 43-44, 47-48, 52; C. asteromphalus
44: C. curvatulus 43; C. marginatus 44, 52; C.
oculus-iridis 44; C. symbolophorus 47; C. tem-
perei 48; C. undulosus 48
Cosmiodiscis 45; C. insignis 45; C. intersectus 45
Cosmiodiscus 27, 34, 37-39, 45, 56, 60; C. insignis
27, 34, 37-39, 45, 56, 60
Costa Rica 65-69, 71, 73, 75, 77-79, 153, 160, 169,
175, 186, 198-201, 204
Cottoclinus gen. nov. 155-160; C. canops sp. nov.
155-160
Cricetulus 213
Crotalinae 407, 440, 450
Crotalus scutellatus 407
Cryptobranchia 170, 204, 205
Ctenocella 261
Curimagua bayano 356
Cyclotella insolita 246, 253
Cymatosira debyi 45, 54, 56
Cymatotheca weissflogii 38, 48
Cymbella coamoensis 247, 253
Cyrtodactylus 463-465, 467-469, 471, 473-479,
481-482, 484-489, 491-497; C. aaroni 493: C.
abrae 493; C. adleri 492, 493; C. aequalis sp.
nov. 467, 485, 489, 491-492, 494, 495: C. agusa-
nensis 493; C. albofasciatus 492; C. angularis
492, 493: C. annandalei sp. nov. 465, 467-469,
481, 492-493, 495: C. annulatus 492: C.
ayeyarwadyensis sp. nov. 467, 469-471,
473-475, 478, 482, 492-493, 495, 496: C. baluen-
sis 493; C. brevidactylus 464, 492, 495, 496: C.
brevipalmatus 492: C. cavernicolus 492: C.
chrysopylos sp. nov. 467, 485-489, 492, 494, 495:
C. collegalensis 492; C. condorensis 492-493; C.
consobrinoides 463-464, 468469, 481, 492-493,
495: C. consobrinus 493-495: C. darmanavillei
493; C. deccanensis 492: C. derongo 493: C. elok
492-493; C. feae 464, 493-494: C. fraenatus
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
492-493: C. fumosus 492; C. gansi sp. nov. 467,
475-478, 492-493, 495; C. gubernatoris 493; C.
ingeri 492; C. interdigitalis 493; C. intermedius
492-494; C. irianjayaensis 492-493; C. irregu-
laris 492-493; C. jarujini 493; C. jellesmae 492;
C. jeyporensis 492; C. khasiensis 463-464, 469,
474-475, 478, 486, 492-493, 495; C. laevigatus
492: C. lateralis 492-493; C. loriae 493; C. louisi-
adensis 493; C. lousiadensis 493; C. malayanus
492, 493; C. malcolmsmithi 493; C. marmoratus
492: C. matsuti 492-493; C. mimikanus 493; C.
nebulosus 492; C. novaeguineae 493; C. oldhami
464, 492-494: C. papilionoides 493; C. papuensis
492: C. paradoxus 492; C. peguensis 464, 489,
492-495; C. philippinicus 492; C. pubisulcus 492;
C. pulchellus 464, 492; C. quadrivirgatus
492-493: C. redimiculus 493; C. rubidus 464, 492,
494: C. russelli sp. nov. 467, 482, 484-485,
492-495; C. sadleiri 492: C. sermowaiensis 492;
C. slowinskti 463-464, 469, 479, 485, 493, 495: C.
sumonthai 492-493; C. sworderi 493; C. tioma-
nensis 493; C. (G) triedrus 492: C. tuberculatus
493; C. variegatus 464, 493, 495; C. wakeorum
sp. nov. 467, 479, 481-482, 485, 493- 495; C.
wetariensis 493; C. yakhuna 492; C. yoshii 492
Cybaeus 544; C. potanini 544
D
Daboia 411, 413, 419, 439-440, 453-454; D. elegans
439; D. russelii 411, 413, 419, 439-440, 450,
453-454; D. r. formosensis 440; D. r siamensis
439
Dendrodoris 170, 204; D. grandiflora 170
Dendronotacea 256, 262, 302, 325, 330
Dialommus 155, 158-160; D. fuscus 155, 158-160;
D. macrocephalus 155, 158-160
Delphineis 38-40, 45, 56, 58; D. angustata 45, 56; D.
ischaboensis 45; D. karstenii 45; D. sachalinensis
45, 56; D. simonsenii 38, 45, 58; D. surirella 40,
45
Delphines [sic] 34, 37; D. . simonsenii 34, 37
Denticula kamtschatica 46
Denticulopsis kamtschatica 46
Detonula 34, 37-38, 40, 45, 50, 58; D. confervacea
34, 37-38, 45, 50, 58
Diatoms 27-31, 34, 37-43, 245-249, 251, 253-254
Diaulula 65-66, 71-72, 74-78; D. aurila 65-66,
75-78; D. greeleyi 65-66, 71-72, 74-75, 78; D.
punctuolata 78; D. sandiegensis 78
Dicladia 45, 52; D. capreola 45; D. capreolus 45, 52;
D. pylea 45
INDEX TO VOLUME 54
Dinodon septentrionalis 409
Diploneis 247, 253; D. gravelleana 247, 253; D.
smithii 45; D. smithii var. adversa 247, 253
Discodorididae 169-170, 205-206, 394
Discodoris 65-66, 75, 78—79,170, 202-204, 206; D.
aurila 65-66, 75, 78-79; D. boholiensis 170,
202-203
Dispholidus typus 407
Distephanus 48; D. crux 48; D. speculum 48; D.
speculum pentagonus 48; D. speculum speculum
48
Disteira 413, 418, 429-430, 451; D. doliata 429; D.
nigrocincta 418, 429; D. schistosa 430
Dolicamphilius [sic] 90, 94, 108, 125, 129; D. longi-
ceps 90, 94
Dolichamphilius gen. nov. 82, 85, 87, 89, 100-101,
110-111; D. brieni 89, 100, 110, 129; D. longiceps
sp. nov. 82, 87, 94, 100, 110, 129
Doradidae 109
Doratonotus 361
Doridacea 65, 79
Doris 167, 170, 174, 190, 207; D. grandiflora 170; D.
grandifloriger 170; D. nodulosa 174, 190; D.
novae-zelaniae 190; D. pustulosa 190
Doumea 83, 87-88
Doumeinae 83, 87—88, 109-110
Draconarius 499, 500, 507, 510, 537, 561, 564, 575,
576, 662; D. absentis sp. nov. 499, 509, 514,
516-517, 585, 647; D. acidentatus 500, 509-510,
518, 586, 645, 647; D. adligansus 500, 509, 511,
518, 528, 586, 645, 647; D. agrestis sp. nov. 499,
509, 517, 519, 532, 587, 647; D. altissimus 500,
SOI IZ S16) S19, 5215531, 5345551, 6473D:
amygdaliformis 500, 509, 515, 520, 647; D. arcu-
atus 509, 513, 520, 525, 588, 647; D. argenteus
500, 509, 515, 521, 540, 588, 647; D. aspinatus
508, 513, 516, 519, 521, 555, 589, 648; D. bacca-
tus 500, 510, 540; D. baronii 509, 517, 521, 552,
590, 648; D. baxiantaiensis sp. nov. 499, 508,
513, 522, 544, 590, 645, 648; D. bituberculatus
500, 509, 515, 522, 591, 648; D. brunneus 500,
509, 515, 523, 529, 541, 545, 648; D. calcari-
FOMMISHS08, «512, 516, 323) 925; 527, 595; 7962,
591, 645, 648: D. capitulatus sp. nov. 499, 509,
515, 524, 529, 545, 592, 648; D. carinatus 500,
509, 513, 515, 524, 592, 649; D. chaigiaoensis
500, 509, 515, 525, 593, 645, 648; D. cheni 509,
513, 520, 525, 593, 649; D. colubrinus 508, 512,
316, 523; 525; D: coreanus 508; 512, 517; 526,
543, 555, 649; D. curiosus sp. nov. 499, 510, 514,
516, 526, 532, 595, 649; D. davidi 508, 512, 523,
527, 555, 596, 649; D. denisi 500, 510-511, 519,
693
528, 596, 649; D. digitusiformis 500, 510-511,
516, 528, 597, 645, 649; D. disgregus sp. nov.
499, 510, 515-516, 523, 528, 598, 649; D. dissitus
sp. nov. 499, 510, 515, 529, 598, 650; D. dubius
sp. nov. 499, 510, 515-516, 530, 535, 542, 548,
599, 650; D. episomos sp. nov. 499, 509, 512,
530, 599, 650; D. everesti 500, 510, 517, 531, 549,
650; D. funiushanensis 500, 508, 512, 517, 532,
535, 650; D. griswoldi sp. nov. 499, 510, 514,
517, 519, 527, 531, 600, 651; D. gurkha 509, 511,
532, 549, 600, 650; D. gyriniformis 500, 508, 513,
533, 650; D. hangzhouensis 500, 510-510, 534,
601, 651; D. haopingensis sp. nov. 499, 508, 514,
533, 539, 601, 643, 650; D. himalayaensis 500,
SO9F Sl 516; 520)9319 934559425951 Da nut
500, 508, 512, 532, 534, 651; D. huizhunesis 500,
507, 509, 511, 516, 535, 541, 550, 651; D. incer-
tus sp. nov. 499, 510, 535, 602, 651; D. infulatus
510-511, 536, 651; D. jiangyongensis 500, 510,
513, 536, 603, 643, 651; D. labiatus 508, 513,
536, 537, 603, 652; D. linxiaensis sp. nov. 499,
510, 516, 537, 540, 604, 652; D. linzhiensis 500,
508, 512, 537, 546, 555, 652; D. lutulentus 500,
508, 513, 517, 538, 605-606, 652; D. magniceps
500, 510, 514, 538, 607, 652; D. molluscus 508,
514, 533, 539, 607, 643, 652; D. nanyuensis 500,
510, 512, 539, 607, 652; D. neixiangensis 500,
510-511, 516, 526, 537, 540, 608, 652; D. nudu-
lus sp. nov. 499, 510, 515, 521, 540, 608, 653; D.
ornatus 500, 509, 516, 535, 541, 551, 609, 643,
653; D. parabrunneus sp. nov. 499, 510, 514,
541, 545, 609, 643, 653; D. paraterebratus sp.
nov. 499, 510, 513, 542, 609, 653; D. parawudan-
gensis 499, 508, 554; D. patellabifidus sp. nov.
499, 510, 515-516, 530, 534, 542, 548, 610, 654;
D. penicillatus 500, 510, 511, 517, 543, 611, 653;
D. pervicax 500, 510, 513, 543, 546, 653; D. picta
500, 508, 512, 544, 555, 653; D. potanini 510,
513, 522, 544, 611, 653; D. pseudobrunneus sp.
nov. 499, 510, 514, 541, 544, 612, 654; D.
pseudocapitulatus sp. nov. 499, 510, 515, 524,
529, 545, 612, 655; D. pseudowuermlii sp. nov.
499, 510, 514, 546, 555, 612, 654; D. gingzangen-
sis 500, 508, 512, 537, 546, 555, 654; D. quadra-
tus 500, 510, 514, 544, 546, 613, 654; D. rotundus
sp. nov. 499, 510-511, 547, 613, 654; D. rufulus
500, 509, 514-515, 547, 614, 643, 654; D.
schenkeli 510, 511, 548, 615, 654; D. shuang-
paiensis 645; D. simplicidens sp. nov. 499, 510,
515, 530, 542, 548, 615, 655; D. singulatus 510,
511, 517, 531, 533, 549, 616, 655; D. sinualis 500,
508, 538; D. stemmleri 508, 513, 549, 550, 552,
694
556, 617, 655; D. streptus 500, 510, 514, 550-551,
553, 655; D. striolatus 508, 515, 550, 617, 655; D.
strophadatus 500, 509, 511, 535, 550, 655; D. sub-
titanus 500, 509, 512, 520, 531, 534, 551, 655; D.
syzygiatus 500, 510, 514, 516, 550-551, 553, 656;
D. terebratus 500, 509, 514, 516, 541-542, 551,
618, 643, 656; D. tibetensis sp. nov. 499, 510,
517, 552, 619, 656; D. trifasciatus 508, 513, 517,
552, 656; D. tryblionatus 500, 510, 514, 516, 553,
656; D. uncinatus 500, 510, 516, 553, 620, 643,
656; D. venustus 507-508, 512, 517, 554-555,
576, 656; D. wenzhouensis 508, 508, 513, 537,
554, 620, 656; D. wudangensis 499, 508, 512, 517,
Ses 2 6993" 5381994451554) 1621) 6572, D.
wuermlit 508, 513, 538, 546, 555, 622, 657; D.
yadongensis 500, 508, 513, 549, 556, 657; D.
yichengensis sp. nov. 499, 510-511, 556, 622,
657; D. yosiianus 508, 512, 556, 657
Drejerella 373
E
Easter Island: See Pacific Ocean
Eastern Pacific 169, 198, 292,
Panama 437
Echidna nocturna 159
Egypt 321; Suez Canal 303, 321, 330
Elapidae 407, 413, 421, 450-452, 497; Elapinae 421;
Hydrophiinae 407, 427
Elaps 421, 424, 427; E. bungaroides 421; E. macclel-
landti 427; E. maculiceps 424
Elephas 213-214, 221
Ellisella 261
Ellisellidae 255—256, 260-262
Enhydrina 411, 413, 416, 429-430, 436; E.
429; E. schistosa 411, 413, 416, 429-430
Enhydris 407
Epeirotypus 359
Epibulus 361
Equus 217, 219, 220
Erinaceidae 213
Eunotia 247, 253; E. indica var. undulata 247, 253: E.
lunaris var. duolineata 247, 253
Eurocoelotes 574; E. inermis 575
F
Facelinidae 161, 167
Femoracoelotes 499-500, 503, 557, 576, 623-624,
658; F latus 557-558, 623, 658; F. platnicki 557,
624, 658
Fiji: See South Pacific
300, 381; Gulf of
zweifeli
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
Flabellina 328
Fordonia 407
Fragilariopsis 38, 45, 54; F- cylindrus 38, 45, 54; F
oceanica 38, 45, 54
G
Galapagos Islands 155-156, 437; Espanola (Hood
Island) 155-156, 159
Gagata 85, 114
Gargamella 204-205
Garra 104
Gastropoda 391-392, 406, 672, 674, 676, 680
Geckoella 464, 492
Geitodoris 169-170, 202-204, 206; G planata 170,
202
Gekkonidae 463, 465, 496-498
Gephryoglanis 82; G. rotundiceps 82
Gomphonema carolinense 247, 253
Goslineria 205
Grammatophora 45; G. angulosa 45
Greece 321
Guatemala 371—373, 377-378, 380; Izabal 377; Petén
378
Guinea 90, 95, 110
Gulf of Siam 431, 435, 438
Gymnodactylus consobrinoides 465
Gyrosigma 247-248, 253; G acuminatum var. angu-
latum 247, 253; G vartipunctatum 247, 253; G
variistriatum 248, 253
H
Halgerda 393-406; H. aurantiomaculata 406; H.
dichromis 393-394, 398, 399, 400; H. elegans
404-405; H. formosa 394, 397, 399, 406; H.
graphica 393-399, 401, 403, 405; H. gunnessi
397, 398; H. iota 403-404; H. johnsonorum
397-398, 404; H. okinawa 393-394, 397, 399,
401-404; H. terramtuentis 397; H. willeyi
393-394, 397-398, 402, 404-405
Hallaxa 168, 328, 330
HAM 663, 665-667, 669-674, 676-680; See also
Hamus
Hamadryas 426; H. elaps 426; H. hannah 426; See
also Ophiophagus
Hamus 665-676, 679-680, 684; See also Hypotheti-
cal taxa
Hawaiian Islands 169, 171-172, 175, 177-178,
183-185, 189, 204-205, 207, 283-284, 286-287,
300, 302, 307-308, 311, 316-317, 329, 361, 406,
437; Oahu 171, 185, 311, 316-317
INDEX TO VOLUME 54
Hemibungarus: H. macclellandi 427; H. m. macclel-
landi 427
Hercotheca mamillaris 45
Himalayas 1, 209, 357, 427, 441, 574-576
Himalcoelotes 500, 521-522, 575; H. brignolii
521-522
Holocene 209, 215, 217, 224
Homeocladia vidovichii var. nodulosa 248, 253
Homiodoris novaezelandiae 190
Homo sapiens 217
Homoiodoris novaezelandiae 190, 207
Hoplodoris 169-207; H. armata 169, 202-203, 206;
H. bifurcata 169, 185-189, 194-195, 197-198,
202-206; H. bramale sp. nov. 169, 186, 190,
198-204, 206; H. desmoparypha 169-170,
174-175, 177-178, 194, 202-206; H. estrelyado
169, 190-191, 194, 202, 204, 206; H. flammea sp.
nov. 169, 186, 194-198, 202-204, 206; H. grandi-
flora 169-170, 172-184, 194, 200-206; H. mauri-
tiana 202, 203, 204; H. nodulosa 169, 185-186,
190-194, 200-204, 206; H. novaezelandiae 169,
190-191, 206
Huxley’s Archetypical Mollusc 665; See also HAM
Hyalodiscus obsoletus 45, 50
Hyalopyxis concava 45-46, 48, 54
Hydrophiinae (See Elapidae)
Aydrophis 411, 415-419, 429-435, 451, 453; H. atri-
ceps 417, 430, 431-432; H. (Aydrophis) atriceps
417; H. caerulescens 418, 431; H. (Aturia)
caerulescens 419: H. cantoris 411, 416-416, 431;
H. (Hydrophis) cantoris 418; H. fasciatus 411,
430-432; H. (Hydrophis) fasciatus 417; H. fascia-
tus atriceps 430; H. flaviceps 431; H. gracilis 411,
415, 417-418, 432; H. (Hydrophis) gracilis 418;
H. nigrocinctus 429; H. obscura 433; H. obscurus
411, 433; H. (Hydrophis) obscurus 418; H. orna-
tus 411, 433; H. o. maresinensis 434; H. o. ocella-
tus 434: H. o. ornatus 434; H. (Aturia) ornatus
418: H. ornatus ornatus 433; H. schistosus 429,
430; H. spiralis 411, 417-418, 434; H.
(Leioselasma) spiralis 418; H. stricticollis 411,
434 435; H. (Aturia) stricticollis 418
Hydropotes 213, 216-217, 219, 222
HAydrus 427, 429, 430, 431, 432, 434; H. caerulescens
431; H. colubrinus 427; H. fasciatus 430, 432; H.
gracilis 432; H. major 429; H. spiralis 434; H.:
valakadyn 429
Hyomacrurus 279-283, 285, 293-294, 296-298; H.
heyningeni 280; H. hyostomus 280, 294, 298
Hypothetical ancestral mollusc 663; See also HAM;
See also Hypothetical taxa
Hypothetical taxa (Genera Hamus and Protohamus):
695
H. barcalolivorum 667, 670, 674, 684; H. barnesi
667, 674, 676, 684; H. barthbrosorum 667, 684;
H. borrapottsorum 667-668, 673, 684; H. bossi
667, 684; H. eastoni 667, 672-673, 675-676, 684:
H. hickmani 667, 684; H. huxleyi 666, 667,
671-672, 684; H. lankesteri 667-668, 670,
672-673, 684; H. meglitschi 667, 674, 684; H.
moolafisherorum 667, 672-673, 679, 684; H. mor-
toni 667-670, 674, 684; H. mortonyongorum
667-668, 670, 674, 676, 684; Hamus (Schinkenus)
naefi 665; H. nicholsoni 667-668, 672-673, 675,
684; H. pelseneeri 665, 667-668, 672-673, 684;
H. pojetarunnegarorum 669, 670, 684; H.
(Jambonus) portmanni 665; H. ravenjohnsonorum
667, 669-670, 674-675, 684; H. russellhunteri
667, 674-676, 684; H. salvinisteinerorum 667,
670, 674, 684; H. seedi 667-670, 684; H. staseki
667, 684; H. swinnertoni 667, 672-676, 684; H.
yongethompsonorum 667-670, 676, 684; Proto-
hamus verrilli 665; P. yongi 666
Fy strix 2223) 22205222 ,6203
I
In Memoriam: Yuri [gorevich Sazonov 299; Joseph
Bruno Slowinski 411
Ictaluridae 110
Iberian Peninsula 382; See also Portugal; See also
Spain
India 1-2, 11, 14-15, 24, 26, 141, 146, 151-153, 419,
421-422, 426-428, 430-433, 435-436, 439-446,
449-451, 453, 463-465, 486, 494, 496-497;
Assam 11, 14, 26, 141, 421, 425-427, 443-446,
486, 495-496; Bengal 421, 425-426, 429-431,
435, 446, 496; Khasi Hills 11, 421, 442; Madras
151; Orissa 435; Palni Hills 151; Sikkim 421, 427,
440, 446
Indian Ocean 167, 279, 281, 284-287, 290, 300-301,
318, 329, 331, 431; Andaman and Nicobar Islands
448, 464; Bay of Bengal 283, 287-288, 428-431,
435; Maldives 284, 287; Mascarene Ridge
287-288, 290-291; Ninety East Ridge 290-291,
295; Saya de Malha Bank 281, 284-286, 290-291;
Socotra 284, 286-287
Indonesia 1, 15, 113, 153, 169, 191, 194, 255, 259,
288-289, 311, 315, 329, 369, 422,.425-426, 428,
430-432, 435, 437, 440-442, 445, 447-448;
Ambon 1; Bali 169, 175, 186, 194, 196-197, 204;
Batjan 369; Coral Sea 191, 361, 364-365, 369:
Java 422, 428, 432, 438, 440; Kalimantan 422,
447; Lomblen Island 440; Mentavay Ridge
287-288; Moluccas 1, 15; Sulawesi (Celebes)
696
288, 294, 296, 435, 445: Sumatra 1, 284, 286-287,
422, 427-428, 432, 440-441, 447-448, 450, 494;
See also Borneo; See also Straits of Malacca
Indo-Pacific 161, 167-168, 204, 207, 260, 262,
283-284, 303, 311-312, 314, 322-323, 328, 330,
361, 381, 405, 406
Iran 1, 151; Seistan and Baluchestan Province 151
J
Jacunia papillosa 315
Japan 25, 34, 37, 39-41, 43, 167, 185, 189, 206, 257,
DS ORD OlN2632655 26142695, 27) 1, 273. 275214.
BOQ 0883169315, 32153230 329513565 3700392.
399, 401-403, 405, 428, 436, 501, 564, 566-569,
575-578; Honshu 315, 399; Izu Peninsula 399,
401; Ryukyu Islands 185, 312-313, 315, 321, 402,
434, 578: Okinawa 167, 185-189, 205, 255-257,
259-261, 263, 265, 267, 269, 271, 273, 275, 277,
302, 312-313, 315, 321, 369, 391, 393-394, 397,
399-406
Jorunna 205
Justicia 371-380; J. angustiflora 378; J. dendropila
sp. nov. 371-376, 378; J. jitotolana 378; J. lin-
denii 378; J. masiaca 378; J. multicaulis 378; J.
nelsonii 378; J. nevlingii 378; J. stellata 375; J.
tabascina 371, 373, 377-378; J. valvata 371, 373,
376-378; J. SWS Basie db
warnockii 378
viridescens
K
Kerilia 411, 415-416, 435; K. jerdoni 411, 415-416,
435
Kinshasa 90, 92, 94, 96-97, 99-100, 104-107, 118
Kryptopterus 85
Kumba 282, 300, 301
Kurixalus 23; K. eiffingeri 24; K. idiootocus 24
Kuronezumia 282, 301
Korea 501, 507, 526, 566-567, 575-577
L
Labrisomidae 155,
Labrisomus 158-159
Laos 425
Lapemis 413, 417, 419, 436, 450, 453; L. curtis 436;
L. curtis hardwickii 436; L. hardwickii 417, 419,
436
Laticauda 407, 411, 413, 416, 424-425, 451: L. colu-
brina 407, 411, 416, 427-428: L. laticaudata 407,
411, 413, 416, 428: L. scutata 427
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
Leiocassis 85
Leptocoelotes 499-500, 558-559, 566, 576, 625-626,
658; L. edentulus 558-559, 625, 658; L. pseudolu-
niformis 558-559, 626, 658
Leptoglaninae subfam. nov. 81—91, 93, 95, 97, 99,
101, 103, 105, 107-113, 115, 117-119, 121—125,
127, 129, 131
Leptoglanis 81-84, 86-87, 89, 91-92, 94-95,
97-100, 107-111, 114, 117, 119-122, 124: L. bre-
vis 81, 98-99; L. flavomaculatus 95, 96; L. man-
devillei 81, 94, 97; L. rotundiceps 81, 94, 98; L.
rotundiceps 84, 98-99; L. bouilloni 81; L. brieni
81, 100; L. camerunensis 81, 83, 94; L. cf. dorae
84, 95; L. dorae 95; L. xenognathus 81-83, 87, 89,
91-92, 100, 107-108, 110-111, 117, 119-122, 124
Lepus 213, 220, 222, 223
Lionurus 282
Longicoelotes 499-500, 559-560, 576, 627, 659; L.
karschi 559-560, 627, 659; L. kulianganus 500,
559-560, 659; L. senkakuensis 500, 559-560, 659
Louteridium 371, 380
Liradiscus ellipticus 46
Lithodesmium 34, 38, 46, 62; L. minusculum 34, 38,
46, 62
Lucigadus 282, 293
Lufengpithecus 221
Lycodon 409; L. fasciatus 409; L. zawi 11, 409
M
Mabuya homolacephala 234
Macaca 217, 219, 220; M. mulatta 219
Macrocystis 310
Macrouridae 279, 282, 300, 301
Macrourus 280-282, 284, 288, 293-294; M. hyosto-
mus 280-282, 293-294; M. microstomus 284
Macrurus 281, 283, 292-294; M. heyningeni 293; M.
macrolophus 283; M. microstomus 281; M.
tenuicauda 281, 292, 294
Madagascar 170-172, 175-176, 181-182, 356, 369
Madeira 382, 391
Malacocephalus 292, 293
Malacoctenus 158-159; M. zonogaster 159
Malapteruridae 109
Malapterurus 85
Malaysia 1, 150-152, 422, 424-425, 427-428,
430-432, 435-436, 440, 442, 445, 447-448, 450,
497, 687; Sabah 1; See also Borneo; See also
Straits of Malacca
Malaysian Peninsula 428, 441
Maldives: See Indian Ocean
Marionia 325-328
INDEX TO VOLUME 54
Marshall Islands: See South Pacific
Mastogloia 248, 253; M. obliqua 248;
sancti-johannis 248, 253
Mataeocephalus 279-299, 301; M. acipenserinus
279, 281, 283-287, 292-293; M. (M.) acipenseri-
nus 283; M. adustus 279-280, 282, 288-289, 293;
M. (M.) adustus 283; M. cristatus sp. nov.
279-280, 285, 290-291, 293, 295; M. (M.) crista-
tus 283; M. hyostomus 279, 293-294; M.
(Hyomacrurus) hyostomus 283, 285, 294; M. kotl-
yari sp. nov. 279, 293-294, 296; M.
(Hyomacrurus) kotlyari 283, 285; M. microstomus
279, 284, 286; M. nigrescens 279, 282, 284-287,
293: M. tenuicauda 279, 282, 286, 292-294, 298:
M. (M.) tenuicauda 283
Maticora 423, 424, 452-453; M. bivirgata 424; M.
intestinalis 424; M. lineata 423; M. maculiceps
424: M. nigrescens 424
Mauritius 169, 171
Maymena 359
Mediterranean 321, 331, 381—382; Strait of Gibraltar
381-382, 388
Megalostoma 371-373, 375-379; M. viridescens
371-372, 377
Megatapirus 213, 223
Melanesia: See South Pacific
Melibaea australis 303
Melibe 302-303, 305, 307, 309-319, 321-325,
327-353, 355; M. australis 302-303, 324, 326,
328; M. bucephala 302-303, 305, 316, 318,
323-324, 329, 333; M. capucina 324, 328; M. dig-
itata sp. nov. 302, 305, 307, 309, 312, 314, 321,
324-326, 329, 332, 334-336; M. engeli 302, 307,
309, 324, 329, 332, 337-339, 340; M. fimbriata
315, 321, 323-324, 331; M. japonica 324; M.
leonina 302, 310, 324, 327-328; M. liltvedi 302,
310-311, 319, 324, 328; M. maugeana 324, 328;
M. megaceras 302, 311-312, 314, 324, 327, 329,
331; M. minuta sp. nov. 302, 312, 314, 324, 326,
332, 341-343; M. mirifica 324; M. ocellata 324,
328; M. papillosa 302, 315-316, 318, 322-324,
344-345; M. pilosa 302, 316, 318, 323-324,
345-347; M. rangi 324; M. rosea 302, 311,
318-319, 324, 328, 348; M. tuberculata sp. nov.
3027507, 309; 312, 314,319) 321, 1324-325;,332;
349-351; M. tuberculosa 329; M. vexillifera 316,
321, 323-324; M. viridis 302-303, 305, 312, 316,
318, 321, 323-324, 327, 329, 352-353
Meliboea 303, 321, 323-324; M. fimbriata 323-324;
M. viridis 321, 323
Melosira 43, 45; M. albicans 45
Meroles knoxi 234
M.
697
México 65, 68, 71, 73-74, 225, 227-230, 300, 310,
371-373, 375, 377-380; Baja California 73, 79;
Chiapas 225, 227-230, 379; Nayarit 71, 73-74,
79; Quintana Roo 373, 375; Sonora 68: Tabasco
373, 378-379; Veracruz 373, 377-378
Microcephalophis 431-432; M. cantoris 431; M. gra-
cilis 432
Micromys 213
Microphocommatidae 359
Microtus 213
Miocene 11, 27, 28, 34, 37, 39-43, 114, 221
Mnierpes 155, 158-160; M. macrocephalus 159
Mnierps [sic] 159
Mnierpini 155, 158, 160
Mochokidae 109-110, 113
Mollusca (molluscs) 65, 79, 161, 167-168, 206-207,
255, 262, 329, 330, 391-393, 406, 663-666, 669,
672, 674, 676-683; See also Huxley’s Archetypi-
cal Mollusc; See also HAM; See also Hypothetical
ancestral mollusc
Mozambique 231, 237, 284, 286-287, 321-322
Muntiacus 212-213, 216-217, 219-221, 223
Muraena lentiginosa 159
Mustela 219
Myanmar: 1-5, 7, 9-19, 21, 23-25, 141-147,
149-153, 209, 357, 407, 409-411, 413-415, 417,
419-455, 457, 459, 461, 463-465, 467, 469-471,
473-479, 481-483, 485-487, 489, 491-497;
Ayeyarwady delta 427, 440, 448, 463, 471, 475;
Ayeyarwady Division 421, 425-426, 430, 433,
440, 448, 470, 475; Bago Division 435, 440; Chin
Hills 1, 10-11, 15, 443, 447, 463, 479, 495; Chin
State 3-4, 9-14, 150-151, 425, 443, 447, 449,
474-476, 479; Indo-Burman Range 1, 11, 469;
Kachin State 14, 151, 410, 421, 423, 425, 427,
439, 441, 443-444, 446, 449, 482, 486; Magway
Division 14, 151, 422, 425-426, 440; Mandalay
Division 11, 14, 422-423, 425-426, 427, 440,
496; Mergui Archipelago 431, 435-436; Mon
State 11, 14, 150, 435, 447-448, 463, 469, 489,
492; Rakhine State 11, 19, 21, 25, 141-142,
150-151, 422, 425, 428, 435, 446, 463, 470-471,
474-475, 479, 482; Rakhine Yoma 11, 19, 21, 25,
141-142, 150-151, 443, 463, 469-471, 475, 479,
482, 495; Sagaing Division 10-11, 13-14,
422-423, 425-426, 440, 446, 463, 465, 469, 479,
482, 486; Ponnyadaung Range 10-11; Shan State
11, 13, 409, 423, 440, 449, 463, 486, 489;
Tanintharyi Division (Tenasserim) 422, 426, 431,
433, 435-436, 447-448, 463, 469, 495; Yangon
Division 422-423, 425, 427, 435, 440, 446
Myospalax 213
698
Myotis 213
Mysmena 359
Mysmenidae 359
Mystus 85, 113-114
N
Naatlo 359
Naemorhedus 217, 219, 223
Naja 409, 411, 414, 424-426, 450-452, 497; N. han-
nah (see also Ophiophagus) 426; N. kaouthia 409,
411, 414, 424 425, 453; N. lutescens 424; N. man-
dalayensis 409, 414, 425, 453; N. naja kaouthia
424-425
Nangra 85, 114
Nautilus 677, 683
Navicula 245, 248-249, 253-254; N. borinquensis
245; N. congerana 248, 253; N. disticha 245; N.
expansa 248, 253; N. glacialis 46; N. guaynaboen-
sis 248, 253; N. howeana 248; N. incomposita
249, 253; N. incomposita var. minor 249, 253; N.
lyra var. irregularis 249, 253; N. mannii 249, 253;
N. notanda 245; N. tubulosa var. rhomboidea 249,
253
Nembrothinae 381, 382,
Neodenticula 27-28, 31, S 4
kamtschatica 27-28, 31, 34, 37-39, 4
koizumii 38
Neogene 27, 31, 34-35, 39-43
Neolithic 210, 217, 224
Neopilina 676-677
Nepal 151, 425, 440-441, 443, 445, 451, 507,
532-533, 574-576
New Caledonia 262, 280, 284,
308-309, 361, 366, 369, 428
New Guinea 430, 434, 436, 450
New Zealand 189-191, 207, 360
Nezumia 282, 293, 296
Niger Basin 83, 86, 95, 110
Nile Basin 109-110
Nitzschia 34, 43, 45-46, 54, 249-250, 253; N. brit-
tonii 249; N. cylindra 45; N. cylindrus 45; N.
extincta 46; N. grunowii 45; N. hemistriata 249,
253; N. kamtschatica 34; N. koizumii 34; N. mira-
marensis 250, 253; N. obtusa 250, 253; N. obtusa
var. lata 250, 253; N. obtusa var. undulata 250,
253; N. ponciensis 250, 253; N. quickiana 250,
253; N. rolandii 34, 46, 54
North Pacific 27—28, 31, 34-35, 37-42, 45
Nophodoris 204-205
Nudibranchia 65, 79, 161, 167-169, 206-207, 255,
262, 330, 381, 391, 393, 405406
281, 297,» 300;
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
O
Occidozyga 21
Ochotona 213
Octocorals 168, 255—256, 259-263
Odontella aurita 46, 52
Ogilbia deroyi 159
Ogulnius 359
Ompok 85
Onchidoris 204—205
Ophiophagus 411, 414, 426, 452-453; O. hannah
411, 414, 426, 453
Opisthobranchia 167, 169, 207, 262, 302, 329-330,
391, 405, 406
Otinodoris 204
Ovis 2S 2193223
Oxycheilinus 361, 362-367, 369-370; O. arenatus
361; O. bimaculatus 361; O. celebicus 361; O.
digrammus 361; O. lineatus sp. nov. 362,
364-366, 369; O. mentalis 361; O. nigromargina-
tus sp. nov. 364-366, 369; O. orientalis 361,
369-370; O. unifasciatus 361
Opephora schwartzii 46
Ovophis 410-411, 419, 440-441, 451, 454; O. monti-
cola 411, 419, 440-441; O. m. convictus 441; O.
m. makazayazaya 442; O. m. zayuensis 442; O. m.
monticola 440-441
Ike
Pacific Ocean 381, 405-406, 495; Easter Island 317;
Palau Islands 169-173, 175, 179-181; See also
Eastern Pacific; See also Hawaiian Islands; See
also Indo-Pacific; See also South Pacific; See also
Western Pacific
Pakistan 43 1-432, 440
Palau Islands: See Pacific Ocean
Panama 26, 65, 68, 75, 77, 153, 160, 281, 292, 356,
437; Canal Zone 68, 75
Pangasiidae 85, 110, 114
Panthera 213, 217, 223
Papua New Guinea 171, 289
Paracoelotes 555, 568, 570, 574-575; P. luctuosus
575; P. spinivulvus 575; P. wuermlii 555
Paralia sulcata 38, 46, 62
Paramphilius 83, 87; P. firestonei 87; P. teugelsi 87
Patu 356, 357—360: P. samoensis 356, 358; P. vitien-
sis 356
Pelamis 419, 437, 451; P. platurus 419, 437
Peltodoris 65-66, 71, 73-74, 79; P. greeleyi 65, 71,
73-75; P. nayarita 65-66, 71, 74-75, 79
Perciformes 155
INDEX TO VOLUME 54
Persian Gulf 430, 432, 434-436, 438
Pharodoris 205
Phenilia 261
Philautus 17, 21-23, 26; P. cherrapunjiae 22-24, P.
romeri 22—24
Philippine Islands: 161-162, 170-171, 175-176,
185-186, 191, 194, 279, 282, 284, 286-288, 294,
296, 298, 300-302, 305, 307-309, 319, 321-322,
324, 329, 340, 426, 430, 435-436; Cebu 171;
Luzon 162, 185, 279, 294, 296, 305, 307, 319,
321; Sumilon 170
Philippine seas 284
Phractura 83, 87
Phyllodesmium 161-168; P. briareum 167; P. cole-
mani 167; P. crypticum 166; P. guamensis 161,
166-167; P. horridum 166; P. hyalinum 166; P. iri-
ometense 166; P. kabiranum 166; P. longicirrum
166; P. macphersonae 166; P. magnum 161, 166;
P. opalescens 166; P. parangatum sp. nov.
161-166; P. pecten 166; P. serratum 166-167
Pimelodontidae 112
Pinnularia 245, 250-253; P. laterittata var. minor
245, 250-251, 253; P. titusiana 252-253; P. viridis
var. subconstricta 252-253
Plagiacanthus 373
Plato 359
Platocoelotes 499-500, 502, 507, 561-564, 576,
628-631, 659; P. kailiensis sp. nov. 500, 561—
562, 630, 659; P. icohamatoides 561, 562-563,
629, 659; P. icohamatus 561-562, 659; P. impletus
561-562, 564, 628, 659; P. lichuanensis 561-563,
631, 659
Platydoris 167, 204-205, 328, 330
Pleistocene 209, 213, 215, 217, 220-221, 223-224,
437
Pleurosigma 46, 252-254; P. angusti-convexum
252-253; P. portoricense 252; P. portoticense [sic]
253; P. strigosum var. incisum 252, 254
Pliocene 27, 30-31, 34, 37, 39-43, 221
Polyceridae 381, 382
Polypedates 21, 23; P. leucomystax 21
Porosira 38, 40, 46, 52, 60, 62; P. glacialis 38, 46, 62;
P. punctata 46, 52, 60
Portugal 382, 383
Praescutata 411, 437-438; P. viperina 411, 438
Pribilof Islands 27, 38-39, 41, 45
Proboscia 41, 46; P. alata 46; P. barboi 46
Propemelibe mirifica 303, 323
Propomelibe mirifica 321, 324
Protobothrops 410-411, 419-420, 442-444, 451,
454; P. jerdonii 411, 420, 442; P. kaulbacki 419,
443; P. mucrosquamatus 411, 419, 442, 444
699
Psammodynastes 407
Psammphiletria gen. nov. 82, 84, 87-89, 101-102,
111; P. delicata sp. nov. 82, 87-88, 102, 110, 130:
P. nasuta 82, 84, 87-88, 101-102, 111, 130
Psammphyletria [sic] 88, 90, 100, 102, 108, 111, 123,
125, 130; P. delicatus 90, 111; P. nasuta 88, 90,
102, 108, 123, 125
Pseudoboa fasciata 421
Pseudoboa fasciatus 421
Pseudocordylus nebulosus 236
Pseudopodosira elegans 48
Pseudopyxilla 46, 62; P. americana 46, 62; P. rossica
46
Pterostichini 238
Pterotheca 47; P. kittoniana 47; P. kittoniana var.
kamtschatica 47
Puerto Rico 245, 247, 249, 251, 253
Pulmonata 225, 230
Pyxidicula zabelinae 46, 50
Quaternary 27, 35, 37, 39, 40, 42
R
Rana 21; R. alticola 21; R. erythracea 21; R. lateralis
21; R. limnocharis 21; R. nigrovittata 21
Rattus 212-213
Red Sea 262, 303, 329, 369-370
Republic du Congo 96-97, 99-100, 104-107; See
also Republique du Congo
Republique Centrafricaine 92, 97, 101-102, 104,
106-107
Republique du Congo 92, 94; See also Republic du
Congo
Reunion 170, 172, 174, 181-182
Rhabdonema japonicum 46
Rhabdophis 407, 452, 454; R. subminiatus 452, 454
Rhacophoridae 16-17, 26
Rhacophorus 23
Rhaphoneis 34, 37, 45-46, 56, 60; R. amphiceros 46;
R. amphiceros var. angularis 47; R. angularis 34,
37, 46, 56, 60; R. sachalinensis 45
Rhinoceros 212, 214, 217, 219, 220; R. cf. sinensis
DNS 220,
Rhinocerotidae 213
Rhizomys 213, 217, 219-220, 222-223
Rhizosolenia 46-47, 56; R. alata 46; R. barboi 46; R.
curvirostris var. inermis 46; R. hebetata 47, 56; R.
hebetata f. hiemalis 47; R. hebetata f: semispina
47; R. setigera 47; R. styliformis 47
700
Roboastra 381-392; R. caboverdensis sp. nov. 381,
383, 387-391; R. europaea 381-386, 388-392; R.
gracilis 381, 389, 392; R. luteolineata 381, 392; R.
rubropapulosa 381; R. tigris 381
Robusticoelotes 576
Russia 501, 566-567, 575-576; Far East Russia
575-576; Siberia 437
S
Saudi Arabia 151
Scelotes 231-237; S. anguineus 231; S. arenicola
235; S. bidigittatus 233; S. bipes 231, 233-235; S.
brevipes 233; S. cafer 231; S. gronovii 231,
235-236; S. guentheri 233; S. kasneri 231-236; S.
sexlineatus 231, 233-235; Scelotes montispectus
sp. nov. 232-236
Schilbeidae 108—109
Sciuridae 213, 223
Sciurotamias 213
Sclerodoris 204, 406
Scorpaena mystes 159
Semiconchula 225-230; S. custepecana 225, 229; S.
breedlovei sp. nov. 225-226, 228-230
Senegal Basin 110
Silicoflagellates 30-31, 41-42, 48
Silurichthys 85
Singapore 425, 427, 447, 448, 450
Sinoadapis 221
Sinomicrurus 411, 414-415, 427, 453; S. macclellan-
di 411, 414-415, 427
Sisoridae 82, 107, 113
Socotra: See Indian Ocean
Sokodara 285, 301; S. johnboborum 285; S. misakia
285
Solomon Islands: See under South Pacific
Somalia 283-284, 286-287
Sonorella 229-230
Soricidae 213
South Africa 168, 231, 233-237, 310-311, 318, 328,
398, 399, 405; Cape Province 231, 234-236, 310,
318; KwaZulu-Natal Province 231; Limpopo
Province 231; Mpumalanga Province 231; Natal
398; Western Cape Province 231, 234-235;
Swaziland 231, 237
South China Sea 284, 286-287, 290-291, 295
South Pacific: 191, 361; Cook Islands 361-362,
364-365; Chesterfield and Bellona Plateau 280,
283, 296-298; Chesterfield Bank 361, 364—365,
369; Fiji 356, 358; Loyalty Islands 280, 283, 298:
Markus-Necker Ridge 290-291, 295; Marshall
Islands 191, 369; Melanesia 428, 432; Pitcairn
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
Islands 361, 365; Polynesia 428; Rarotonga
361-362, 364-365; Sala-y-Gomez Ridge 284,
286-287; Samoa 356, 358; Solomon Islands 191,
464; Tahiti 361, 365; Vanuatu 280, 284, 288-289,
296-298; Wallis and Futuna Islands 283-284,
296; See also New Caledonia
Spain 381, 382, 391-392
Sphagemacrurus 282, 293
Spiricoelotes 499-500, 502, 507, 561, 564-566, 576,
631-632, 645, 660; S. pseudozonatus 500, 564,
565, 631, 660; S. urumensis 564—566, 660; S.
zonatus 564-566, 632, 645, 660
Sri Lanka 145, 151-152, 321,
435-436, 440, 464
Starksia galapagensis 159
Staurodoris pustulata 190
Stauroneis similaris 252, 254
Steatoda 133-139; S. cingulata 138; S. mainlingensis
133-134; S. mainlingoides sp. nov. 133, 135; S.
pardalia sp. nov. 133-136; S. terastiosa 137; S.
tortoisea 133, 135, 138-139
Stegodon 212-214, 221; S. cf. yangyiensis 221; S. ele-
phantoides 221
Stegolophodon yanyiensis 221
Stellarima microtrias 47
323, 431-432,
Stephanogonia hanzawae 47, 50
Stephanopyxis 47; S. turris 47
Straits of Malacca 431-432, 435-436
Sus 213; 217; 219-220; 233
Symphytognatha 356, 359; S. globosa 356, 359; S.
imbulunga 356
Symphytognathid spiders 356, 360
Symphytognathidae 356, 359-360
T
Tabascina 371, 373, 376-378; T. lindenii 378
Tadzhikistan 507, 576
Taiwan 25-26, 409, 425, 432, 434, 436, 440-442,
444, 448-449, 451, 501-503, 537, 557-559, 576,
578; See also China
Taiwania 357
Tajikistan 554
Talpidae 213
Tambja 389-391; T. simplex 390-391
Tanzania 170=172, 175) 231832353305356
Taringa 65-66, 69-71, 79; T. aivica 65-66, 69-71,
79: T. aivica aivica 65, 71; T. aivica timia 65-66,
71, 79; T. telopia 66
Tasmania 437; See also Australia
Tegecoelotes 499-500, 558, 566-569, 575-576,
633-635, 660: T. bicaudatus 566-567; T. cora-
INDEX TO VOLUME 54
sides 566-568, 634, 660; T: ignotus 566-568, 635,
660; T. michikoae 566-567, 569, 660; T: muscica-
pus 566-567, 569, 635, 660; T: secundus 566-567,
633, 660
Tegenaria 520, 551, 567-569, 577-578; T. corasides
568, 578; T. domestica 520; T: muscicapa 569; T.
pagana 551; T. secunda 567
Tethydidae 302, 324-325, 328
Tethys 302, 325, 327-329; T. fimbria 325, 328
Tetracamaphilius [sic] 88; T: pectinatus 88
Tetracamphilius gen. nov. 81, 84, 86-88, 90, 92-93,
101-108, 111, 125, 131-132; T. angustifrons 84,
87, 90, 103, 105-106, 111, 131-132; T. clandesti-
nus sp. nov. 81, 84, 88, 103, 105-106, 110-111,
132; T. notatus 87, 90, 103, 105-107, 132; T-
pectinatus sp. nov. 81, 87-88, 90, 92-93,
102-104, 108, 111, 125, 131
Thailand 11, 13, 15, 24-26, 113-114, 150, 153, 303,
329, 419, 422, 424-432, 440-442, 445, 447-448,
450, 453, 464, 493, 496, 498, 577; Chiang Mai
Province 11, 13; Kanchanaburi Province 493
Thalassophina 413, 417, 437-438; T. viperina 413,
417, 437-438
Thalassophis 413, 438; T. viperina 413, 438
Thalalassiosira 48; T. margaritae 48; T: orientalis 48
Thalassionema 37-38, 47, 54; T. nitzschioides 38, 47,
54; T. schraderi 37
Thalassiosira 27, 34, 37-40, 42, 45-48, 50, 52, 54,
56, 60, 62; T: antiqua 34, 47, 54; T: borealis 48; T.
convexa 34, 37, 47, 56, 60; T: decipiens 47-48; T.
delicata 47; T. dolmatovae 47, 50, 54; T. eccentri-
ca 47, 56, 60; T: gravida 38, 47, 52; T. gravida f.
fossilis 47; T: hyalina 38, 47, 54; T. insigna 45; T.
Jacksonii 34, 47, 60; T: jouseae 34, 37-38, 47, 50;
T. kryophila 38, 47, 52; T. latimarginata 27, 34,
37-39, 47, 50, 60; T. leptopus 38, 47, 56; T: linea-
ta 38, 48; T. manifesta 48, 62; T: marujamica 34,
48, 54; T. nativa 48, 50; T. nidilus 48; T: oestrupii
27, 34, 37-39, 48, 54; T. praeoestrupii 34, 37, 40,
48, 62; T. punctata 46; T. sheshukovae 34, 48, 50;
T. temperei 27, 34, 37-39, 48, 54; T. tertiaria 34,
37, 48; T. trifulta 47; T: undulosa 46, 48, 56; T.
usatschevii 46; T. zabelinae 46
Thalassiothrix 48, 54, 56; T. longissima 48; T. robus-
ta 48, 54, 56
Thelotornis capensis 407
Theridiidae 133, 140
Theridiosomatidae 359
Tomicodon chilensis 159
Tonsilla 499-500, 569-575, 636-642, 645, 661-662;
T. eburniformis 570-571, 636, 661; T. imitata
570-572, 661; T. lyratus 570, 572-573, 640, 645,
701
662; T: makros 500, 570, 573-574, 642, 662: T.
tautispinus 570, 572-573, 640, 662; T. truculenta
569-572, 636-640, 661; T. variegatus 570,
573-574, 641, 662
Trachyglanis 83
Trachyneis aspera var. atomus 252, 254
Trachypithecus 220, 222
Trichomycteridae 112
Trigonocephalus 444-445; T. erythrurus 445: T.
mucrosquamatus 444; T: purpureomaculatus 447
Trimeresurus 409-411, 413, 419-420, 440-449,
451-452, 454; T. albolabris 420, 444-445; T. a.
albolabris 444; T. a. insularis 445; T. a. septentri-
onalis 445; T. erythrurus 411, 413, 420, 445-446;
T. flavoviridis 442; T. jerdonii 442; T. j. bourretti
443; T. j. xanthomelas 443; T. kaulbacki 443; T.
medoensis 419, 446, 450, 452; T: monticola 440;
T. m. monticola 440; T. mucrosquamatus 411, 444;
T. popeiorum 411, 420, 446-447; T. popeorum
446-447; T. purpureomaculatus 413, 419-420,
447-448; T. p. purpureomaculatus 447; T. stej-
negeri 409, 411, 420, 447— 450; T. s. stejnegeri
448; T. s. yunnanensis 449; T. yunnanensis 409,
420, 447-449
Tritonia 255-257, 259-263, 265, 267, 269, 271, 273,
275, 277; T. bollandi sp. nov. 255-256, 259-262;
f olivacea 255, 257; 259) 2627269, 271
Tritoniidae 256, 259-262
Trochosira 45-46, 48; T. concava 45; T. spinosa 46,
48
Tropidoneis van-heurckii var. maxima 252, 254
Turkey 152; Icel Province 152
U
United States 73, 171; California 37-38, 68, 71, 73,
310, 319, 328, 437; Florida 73; South Carolina 73,
79; See also Hawaiian Islands
Urocoras 574-575; U. nicomedis 575; U. phthisicus
S15,
Uropterygius macrocephalus 159
Ursus 212-214, 217, 219-220, 223
Vv
Ventrifossa 282, 293, 301
Verrucella 255, 260-261; V. aurantia 255, 260-261
Verspertilionidae 213
Vietnam 24-26, 190-191, 291, 321, 409, 422, 425,
427, 432, 439, 441, 443-445, 447-449
Viminella 261
Vipera 439; V. daboia 439; V. russelli 439; V. russelli
702 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 54, Index
siamensis 439: V. russellii 439 Z
Viperidae 407, 438, 450
Viperinae 407, 438 Zaireichthys 81-84, 86-90, 92-99, 101, 104, 106,
Viverridae 213 108-113, 122-124, 126-128: Z. camerunensis 83,
Volta Basin 110 86-87, 89-90, 93-94, 110, 123, 127; Z. dorae
89-90, 92-93, 95, 98; Z. cf. dorae 84; Z. flavimac-
ulatus [sic] 92; Z. flavomaculatus 90, 92-93, 95,
WwW 98; Z. flavomarginatus [sic; for flavomaculatus]
95; Z. heterurus 84, 86—87, 90, 93, 96-97, 110,
128; Z. mandevillei 84, 87, 90, 93, 96-97, 106,
111, 123, 127-128: Z. rhodesiensis 82; Z. rotundi-
Wadotes 500, 503, 556, 574, 576; W. yadongensis 556
Western Pacific 167, 198, 207, 279, 290, 321, 329
Wermorella 361 ceps 81-84, 86-87, 89-90, 92-93, 95, 97-98, 108,
Cheilinus arenatus 361 128: Z. zonatus 81-83, 87, 90, 92-94, 98, 104,
110; 1135122) 124" 12621071
X Zanzibar 284, 321, 330, 369
Zostera marina 310
Xanthiopyxis 48; X. globosa 48; X. ovalis 48 Zygolophodon 221
Xanthonychidae 225
Xenochrophis 407
Copyright © 2003 by the California Academy of Sciences
San Francisco, California, U.S.A.
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TABLE OF CONTENTS
Marta Poa, JUAN LUCAS CERVERA, AND TERRENCE M. GOSLINER: The Genus Roboastra Bergh,
1877 (Nudibranchia, Polyceridae, Nembrothinae) in the Atlantic Ocean
SHIREEN J. FAHEY AND TERRENCE M. GOSLINER: Redescription of Halgerda graphica Basedow
and Hedley, 1905, with observations on external morphological variation within selected species
of Halgerda (Mollusca, Nudibranchia)
ALAN E. LEVITON, GUIN O.U. WOGAN, MICHELLE S. KOO, GEORGE R. ZUG, RHONDA S. LUCAS,
AND JENS V. VINDUM: The Dangerously Venomous Snakes of Myanmar: Illustrated Checklist
with Keys (with photographs from life by Hla Tun, Dong Lin, and John Tashjian)
AARON M. Bauer: Descriptions of Seven New Cyrtodactylus (Squamata: Gekkonidae) with a
Key to the Species of Myanmar (Burma)
XIN-PING WANG: Species Revision of the Coelotine Spider Genera Bifidocoelotes, Coronilla,
Draconarius, Femoracoelotes, Leptocoelotes, Longicoelotes, Platocoelotes, Spiricoelotes,
Tegecoelotes, and Tonsilla (Araneae, Amaurobiidae)
DaviD R. LINDBERG AND MICHAEL T. GHISELIN: Fact, Theory and Tradition in the Study of
Molluscan Origins
ACKNOWELEDGMENT OF REVIEWERS FOR VOLUME 54
INDEX TO VOLUME 54
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