Table Of ContentPROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
109(4):695-706. 1996
The freshwater crab fauna (Crustacea: Decapoda: Brachyura) of the
Philippines. IV. On a collection of Parathelphusidae from Bohol
Peter K. L. Ng and Boris Sket
(PKLN) School of Biology, National University of Singapore, Kent Ridge, Singapore 119260,
Republic of Singapore; (BS) Department of Biology, University ofLjubljana,
University ofLjubljana, pp 95, Vecna pot 111, 6111 Ljubljana, Slovenia
—
Abstract. ^Five species offreshwater crabs ofthe genus Sundathelphusa are
recognised from the island ofBohol in the Philippines: S. cavernicola (Takeda,
1983), and four new species, S. boex, S. sottoae, S. urichi and S. vedeniki.
Specimens from Bohol previously referredto S. philippina (von Martens, 1868)
belong to S. boex, and those identified as S. cavernicola belong to two separate
species.
The freshwater crab fauna of the Philip- (1991) reported Archipelothelphusa longi-
pines is one of exceptional diversity, al- pes Balss, 1937, from Bautakay Cave in
though this is not apparent from the number Luzon. A total offour species offreshwater
of described species. In recent years, Ng & crabs of the family Parathelphusidae were
Takeda (1992a, 1992b, 1993a, 1993b) have collected by the present expedition. All are
been involved in a systematic revision of undescribed. Of these, three are apparently
this fauna based on extensive collections troglobitic species while one epigeal spe-
made by staff of the National Science Mu- cies makes occasional forays into caves.
seum (Tokyo) and the National Museum of Only two true freshwater species have
the Philippines (Manila). The revision of been reported from Bohol thus far, Sunda-
the two largest genera, Sundathelphusa thelphusa philippina (von Martens, 1868)
Bott, 1969, and Archipelothelphusa Bott, and Archipelothelphusa cavernicola Take-
1969, is now in progress. da, 1983 (Bott 1970, Takeda 1983). Previ-
In February 1995, the Slovene Caving ous collections, however, are rather poor, so
Association launched two caving expedi- the discovery of new taxa is not unexpect-
tions to Asia: to Guizhou, China (Trontelj ed.
1996, Ng & Trontelj 1996), and the Phil- In view of the good series of specimens
ippines (Sket 1995). Altogether, the expe- available from Bohol, the homogeneity na-
dition team investigated about 30 caves in ture ofthe fauna (all the species seem to be
Pleistocene to Miocene aged limestone. The more closely related to each other than to
Philippine island ofBohol in particular, har- others in the Philippines) and the cavemic-
bours one of the largest continuous karst olous habits of the species, it was felt that
areas in the Philippine archipelago (Balasz it would be useful to document this fauna
1973), and because of this, exploration ef- in a single paper.
forts were centeredthere. Studies ofthecol- The generic system used by Bott (1969,
lections made show that Bohol has a rich 1970) for the Southeast Asian fauna is
freshwater crab fauna. problematic. Sundathelphusa Bott, 1969,
Few cavemicolous crabs are known from and Archipelothelphusa Bott, 1969, are so
the Philippines. Takeda (1983) described a close that there seems to be no good reason
new troglobitic species,Archipelothelphusa for separating them. As both Sundathelphu-
cavernicola, from caves in Bohol, and Ng sa and Archipelothelphusa were described
696 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OFWASHINGTON
in the same paper (Bott, 1969), Sundathel- lower part, Cugon, Jagna, Bohol, leg. B.
phusa Bott, 1969, is herein regarded as hav- Sket, Feb 1995—.
ing seniority over Archipelothelphusa Bott, Description. Dorsal surface ofcarapace
1969. In any case, the name Sundathelphu- gently convex; anterolateral regions rugose;
sa Bott, 1969, appears before Archipeloth- posterolateral regions covered with oblique
elphusa Bott, 1969, in Bott's (1969) paper. striae; cervical grooves deep; epigastric
The abbreviations Gl and G2 are used cristae low, rugose, not confluent with low
for the male first and second pleopods re- postorbital cristae; postorbital cristae inter-
spectively. All measurements are indicated rupted medially by cervical groove, not
as carapace width x carapace length. The reaching epibranchial tooth. Frontal median
terminology used follows that by Ng triangle well defined, with dorsal and lateral
(1988). Specimens are deposited in the Na- margins cristate; dorsal ridge not fused with
tional Museum of the Philippines, Manila lateral margins. Anterolateral margin dis-
(NMCR); National Science Museum, To- tinctly convex, smooth; epibranchial tooth
kyo (NSMT); Department of Biology, Uni- low to very low, separated from external
versity of Ljubljana (ULB); and Zoological orbital angle by small notch; posterolateral
Reference Collection, School of Biology, margins gently converging towards poste-
National University of Singapore (ZRC). rior carapace margin. Ocular peduncle and
cornea well developed; eye occupying al-
Systematic Account most entire orbit. Carpus of chelipeds with
surface smooth; inner margin with 1 large
Family Parathelphusidae Alcock, 1910 sublamelliform tooth and 1 smaller tooth.
Genus Sundathelphusa Bott, 1969 Ambulatory legs relatively short; meri of
Sundathelphusa boex, new species third and fourth legs about 2.7 times longer
Figs. lA, 2a-f, 4a-f than broad; dactylus ofthird and fourth legs
— subequal or shorterthan propodus. Male ab-
? Telphusa leschenaulti. Burger, 1894:2
(part) (not Thelpheusa leschenaudii H. domen with slender segment 6, about 1.1
times longer than broad. Gl slender, gently
Milne Edwards, 1834, misspelling by
curved outwards; terminal segment slender,
Burger, 1894).
— conical, about 0.4 times length of subter-
Material examined. Holotype male minal segment. G2 about 1.0 times length
(37.4 X 30.9 mm) (NMCR), small stream ofGl; distal segment well developed, about
near Sierra Bulliones, about 9°42'N, 0.5 times leng—th of basal segment.
124°20'E, about 5 km northwest of Jagna, Discussion. Burger (1894) reported a
Bohol, leg. H. Morioka, 22 Jul 1985. Para- large 50 X 41 mm female from Bohol as
type male (34.2 X 27.8 mm), paratype fe- Telphusa leschenaulti, but his specimen is
male (almost mature) (26.8 X 22.4 mm) most likely S. boex instead. Telpheusa les-
(ZRC 1996.1550-1551), Castigio Cave, chenaudii H. Milne Edwards, 1834 (type
Batuan, Bohol, leg. B. Sket, Feb 1995. locality Pondicherry, India) (spelling ofge-
Paratype male (41.0 X 33.0 mm) (ZRC nus name erroneous) is now regarded as a
1996.1549), Batuan, Bohol, leg. B. Sket, junior synonym of Oziothelphusa senex
Feb 1995. 1 male (NMCR), 1 male (ULB), (Fabricius, 1798) (see Bott 1970:100). The
1 male (NMCR), brook, Pahangong Talon, specific epithet of T. leschenaudii is often
Ginguyuran, Bohol, leg. B. Sket, Feb 1995. spelled "leschenaulti" (changed by H.
1 male (34.6 X 28.8 mm) (NMCR), spring Milne Edwards, 1853) but under current
in Batuan, Bohol, leg. B. Sket, Feb 1995. 1 rules (ICZN 1985), the original speUing
juvenile (NMCR), Capiro Spring, Batuan, must be preserved.
Bohol, leg. B. Sket, Feb 1995. 1 young Sundathelphusaphilippina (von Martens,
male (10.2 by 8.7 mm) (ULB), Carmulaon, 1868) somewhat resembles S. boex, but S.
VOLUME 109, NUMBER4 697
Fig. 1. Dorsal views ofnewSundathelphusa species. A,S. boex, holotype male (37.4 X 30.9 mm)(NMCR);
B, S. sottoae, holotypemale (17.3 X 13.4mm) (NSMT-Cr8938); C, S. urichi, holotypemale (36.6 X 27.9 mm)
(NMCR); D, S. vedeniki, holotype male (28.2 by 22.3 mm) (NMCR).
philippina differs markedly in having a acted as the Philippine component of the
more inflated and proportionately broader expedition. The name is usedhere as anoun
carapace, as well as the differently struc- in apposition.
tured anterolateral margin and Gl. The first
author has examined the types of Sunda- Sundathelphusa cavernicola
thelphusaphilippina in the Berlin Museum. (Takeda, 1983)
Sundathelphusa philippina is known for Fig. 4g
certain only from the islands ofLeyte, Cebu
Archipelothelphusa cavernicola Takeda,
and Samar (PKLN, pers. obs.).
1983:169 (part).
Sundathelphusa boex was found in sur- —
face waters as well as in caves. Both caves Material examined. Holotype female
where the crabs were found are rich in or- (25.7 X 21.0 mm) (NSMT-Cr 8937), mud-
ganic nutrients. Castigio Cave is inside a dy bottom on subterranean stream, 20-30
mogote ("chocolate hill") and has a very cm deep, about 300 m from entrance to east
high amount of plant debris. Carmulaon branch of Quinapon-an Cave, Antequera,
Cave is a vertical cave receiving waters 09°49'38"N, 123°54'10"E, Bohol, leg. S. I.
from nearby rice fields. All specimens are Ueno, 4 Mar 1—983.
usually pigmented. The eyestalks of adults Description. Dorsal surface ofcarapace
are subequally thick both distally and prox- gently convex; anterolateral regions gently
imally, while in juveniles the distal part is rugose; posterolateral regions covered with
proportionatel—y broader. oblique striae; cervical grooves distinct but
Etymology. The species name is de- relatively shallow; epigastric cristae low,
rived from the acronym B.O.E.X. (Bohol rugose, not confluent with low postorbital
Outdoor Explorers Club), whose members cristae; postorbital cristae interrupted me-
698 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OF WASHINGTON
dially by cervical groove, not reaching epi- ferred to S. sottoae, new species (see Dis-
branchial tooth. Frontal median triangle cussion for S. sottoae). Unfortunately, no
poorly defined; lateral margins cristate, dor- males of S. cavernicola are known.
sal margin not cristate or meeting lateral
margins. Anterolateral margin distinctly Sundathelphusa sottoae,
convex, smooth; epibranchial tooth distinct, new species
separated from external orbital angle by Figs. IB, 2g-k, 4h-l
distinct notch; posterolateral margins gently
Archipelothelphusa cavernicola Takeda,
converging towards posterior carapace mar-
1983:169 (part).
gin. Ocular peduncle reduced, cornea —
strongly reduced; eye occupying about half Material examined. Holotype male
of orbit. Surface of chelipedal carpus ru- (17.3 X 13.4 mm) (NSMT-Cr 8938), in
m
gose; inner margin with one large subla- shallow subterranean water, about 50
melliform tooth and one denticle. Ambula- from entrance to Ughob Cave, about 2 km
tory legs relatively long; meri of third and southwest of Batuan, 09°46'45"N, 124°
fourth legs about 5.2 and 3.8 times longer 07'54"E, Bohol, leg. S. I. Ueno, 28 Feb
than broad respectively; dactylus of third 1983. Paratype female (26.0 X 21.0 mm)
and fourth legs distinctly longer than prop- (ZRC 1996.1553), Bonugan, Batuan, Bo-
odus. Male abdomen, Gl and G2 not hol, leg. B. Sket, Feb 1995. Paratype female
known. — (28.0 X 25.0 mm) (NMCR), upper part of
Discussion. This species is here trans- cave, Bonugan Cave, Batuan, Bohol, leg. B.
ferred to Sundathelphusa in line with our Sket, Feb 1995. 1 male (14.8 X 12.5 mm)
proposed synonymy of Archipelothelphusa (ULB), Kalumpan, Behind-the-Clouds, Ba-
and Sundathelphusa. Takeda (1983) de- tuan, Bohol, leg. B. Sket, Feb 1995. 1 male
scribed this species from one large female (17.4 X 13.8 mm) (ZRC 1996.1548), open
and one small male from different caves in well, Batuan, B—ohol, leg. B. Sket, Feb 1995.
Bohol. The holotype female possesses Description. Dorsal surface ofcarapace
strongly reduced eyes, the cornea being gently convex; anterolateral regions gently
highly degenerated. The paratype male, rugose; posterolateral regions covered with
however, possessed a larger cornea and oblique striae; cervical grooves distinct but
more developed eyes. Takeda (pers. comm.) relatively shallow; epigastric cristae low,
has also expressed doubts as to the conspe- rugose, barely confluent with low postor-
cificity ofthe two specimens. The localities bital cristae; postorbital cristae interrupted
where the two types were collected (Qui- medially by cervical groove, not reaching
napon-an and Ughob Caves) are on differ- epibranchial tooth. Frontal median triangle
ent parts of the island. poorly defined; lateral margins cristate, dor-
The good series of specimens from the sal margin not cristate and not meeting lat-
Batuan area (where Ughob Cave is located) eral margins. Anterolateral margin distinct-
confirms the suspicion that the specimens ly convex, smooth; epibranchial tooth dis-
which have been previously referred to tinct, separated from external orbital angle
'''Archipelothelphusa cavernicola''' (fide by distinct notch; posterolateral margins
Takeda, 1983) actually belong to two dis- gently converging towards posterior cara-
tinct species, easily separated by the degree pace margin. Ocular peduncle and cornea
%
of degeneration of the eyes and cornea, as reduced; eye occupying about of orbit.
well as proportions of the third ambulatory Surface of chelipedal carpus rugose; inner
merus. One species, S. cavernicola, is rep- margin with 1 large sublamelliform tooth
resented only by a single female specimen and one denticle. Ambulatory legs relative-
(the holotype) whilst the others (including ly long; meri of third and fourth legs 3.4-
the paratype male ofA. cavernicola) is re- 3.5 and 3.4-3.6 times longer than broad re-
VOLUME 109, NUMBER4 699
spectively; dactylus ofthird and fourth legs termine. In general carapace and leg mor-
distinctly longer than propodus. Male ab- phology, the Kalumpan and Batuan speci-
domen with segment 6 about 1.1 times lon- mens agree best with the holotype male of
ger than broad. Gl slender, gently curved S. sottoae. They nevertheless differ in hav-
outwards; terminal segment slender, distal ing the eye occupying more of the orbit
part especially slim, about 0.3 times length (0.74-0.75 vs. 0.63-0.66), the cornea is
of subterminal segment. G2 about 1.1 times proportionately larger (relative to the whole
length of Gl; distal segment well devel- eye) (0.35-0.37 vs. 0.25-0.31), the tooth on
oped, about 0.5 times length of basal seg- the inner angle ofthe chelipedal carpus pro-
ment. — portionately shorter, the length of the last
Discussion. Sundathelphusa sottoae, ambulatory dactylus being about 7.1 times
new species, is very similar to S. caverni- longer than the maximum width (excluding
cola, both species sharing a high carapace spines) (vs. about 10 times), andthe median
in which the anterolateral margins are gent- part of the subterminal segment of the Gl
ly convex, lateral regions have distinct stri- being more slender, with the tip of the ter-
ae, reduced eyes, as well as elongate am- minal segment gently but distinctly up-
bulatory dactyli. They differ, however, turned (vs. straight). These differences sug-
markedly in the condition of the eyes, gest the possibility that these specimens
which in S. cavernicola are far more re- represent two species. However, until larger
duced than those in S. sottoae. Also distinct specimens from Batuan and Kalumpan are
are the proportions of the ambulatory legs, collected, this cannot be ascertained.
with the meri of S. cavernicola relatively Sundathelphusa sottoae was found in a
longer, especially that of the third ambula- number of rather diverse habitats, all in the
tory leg (length/width ratio = 5.2 and 3.4- Batuan region in the center of Bohol. Pig-
mented specimens originate from a surface
3.5 respectively).
spring pool. The specimen (male, 17.4 X
In his description ofS. cavernicola, Tak-
eda (1983: 172) stated that the paratype 13.8 mm, ZRC) from a garden well in the
male ofS. cavernicola agrees "... with the middle of Batuan is pale, while those from
within caves (Ughob, Bonugan and Kal-
holotype female in the general formation of
umpan: NSMT-Cr 8938, ZRC, NMCR,
the carapace, chelipeds and ambulatory
ULB) are poorly pigmented. Bonugan Cave
legs, but the cornea is not as strongly re-
duced and the eyestalk is slightly movable is one of a chain of caves along the same
brook. The substrate of this stream is only
and occupies about two-thirds the longer
moderately enriched with organic matter
axis of the orbit. ..." Our specimens from
and the surface fauna (including Anura) is
the Batuan area, where the paratype male
present only near the entrances. In Kalum-
of S. cavernicola was collected, has al- pan, the crab was found among stones in
lowed us to ascertain thatTakeda's paratype
an illuminated siphon pool which had no
hmearlee cahcotusaelnlyasreptrheesehnotlsotSy.pesotftooraet,hisanndeiws dirEetcytmcoolnongeyc.t—ioTnhweitshpeacniyessuisrfnaacemesdtreaafmte.r
species.
Prof. Dr. Filipina Sotto of the Marine Bi-
Two of the larger females from Bonugan ology section at the University of San Car-
Cave (ZRC, NMCR) agree very well with los, Cebu City, whose help with logistics
the holotype male for S. sottoae and con- contributed substantially to the successful
firms the usefulness of the diagnostic char-
expedition.
acters used to separate this species from S.
cavernicola. The identities of the two Sundathelphusa urichi, new species
smaller males (14.8 X 12.5 mm, ULB, and Figs. IC, 3a-f—, 4m-p
17.4 X 13.8 mm, ZRC) from Kalumpan and Material examined. Holotype male
Batuan respectively are more difficultto de- (36.6 X 27.9 mm) (NMCR), Quilas Cave,
700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Nueva Vida Norte, Batuan, Bohol, leg. B. anterolateral margins. In physiognomy, S.
Sket, Feb 1995. Paratype male, paratype fe- urichi is closer to S. vedeniki, but the car-
male (ULB), 2 paratype males (28.0 X 20.0 apace ofS. urichi is distinctly more swollen
mm [crushed], 21.6 X 15.5 mm), paratype than S. vedeniki.
female (32.6 X 25.0 mm) (ZRC 1996. Sundathelphusa urichi was found only in
1554-1556), same data as holotype. 1 ju- Quilas Cave, which is in the central plateau
venile (NMCR), Quilas Cave, Nueva Vida east of Batuan. The cave is part of a long
Norte, Batuan, Bohol, leg. B. Sket, Feb chain of chambers with large pools. They
1995. — do not seem to be connected directly to any
Description. Dorsal surface ofcarapace permanent surface stream butrather, are fed
strongly convex; anterolateral regions diffusely or by periodical inputs from the
smooth; posterolateral regions with very surface. However, the presence ofpigment-
low oblique striae; cervical grooves deep; ed catfish (Clariidae: Clarias sp.) in the
epigastric cristae very low, rugose, not con- cave suggests that a hidden connection to
fluent with very low postorbital cristae; surface waters might exist. The rich organic
postorbital cristae interrupted medially by matter present, mostly detritus from the sur-
cervical groove, not reaching epibranchial face, supports a rich population of shrimps
tooth. Frontal median triangle poorly de- (Decapoda: Atyidae), with amphipods {Er-
fined; lateral margins cristate, dorsal margin iopisa sp.) being less numerous. Many
not cristate or meeting lateral margins. An- specimens ofthis amphibious crab wereob-
terolateral margin strongly convex, smooth; served. —
epibranchial tooth very low, separated from Etymology. The present species hon-
external orbital angle by faint but distinct ours Dr. Peter Urich, a "caver" and socio-
notch; posterolateral margins strongly con- geographer, now at the Waikato University,
verging towards posterior carapace margin. New Zealand. An expert on Boholano so-
Ocular peduncle and cornea slightly re- ciety and nature, he efficiently took care of
duced; eye occupying about 3/4 of orbit. the expedition group.
Surface of chelipedal carpus smooth; inner
margin with 1 large sublamelliform tooth Sundathelphusa vedeniki,
and 1 denticle. Ambulatory legs relatively new species
long; meri ofthird and fourth legs about 3.7
Figs. ID, 3g-l, 4q, r
and 3.5 times longer than broad respective- —
ly; dactylus of third leg subequal to length Material examined. Holotype male
of propodus; dactylus of fourth leg longer (28.2 X 22.3 mm) (NMCR), Boho sa Bi-
than propodus. Male abdomen with slender kahan, Bikahan, Antequera, Bohol, leg. B.
segment 6, medially constricted, about 1.2 Sket, Feb 1995. Paratype male (33.9 X 26.5
times longer than broad. Gl slender, gently mm) (ZRC 1996.1552), same data as ho-
curved outwards; terminal segment slender, lotype. —
conical, about 0.4 times length of subter- Description. Dorsal surface ofcarapace
minal segment. G2 about 1.2 times length distinctly convex; anterolateral regions ru-
ofGl; distal segment well developed, about gose; posterolateral regions with distinct
0.4 times lengt—h of basal segment. oblique striae; cervical grooves deep; epi-
Discussion. Sundathelphusa urichi, gastric cristae low, rugose, not confluent
new species, differs markedly from S. sot- with low postorbital cristae; postorbital
toae, new species (which occurs in the cristae interrupted medially by cervical
same area) in having distinctly proportion- groove, not reaching epibranchial tooth.
ately shorter and more falcate ambulatory Frontal median triangle not well defined;
dactyli (straighter in S. sottoae), a distinctly lateral margins cristate, dorsal margin
more swollen carapace and more convex weakly cristate and not meeting lateral mar-
VOLUME 109, NUMBER4 701
—
gins. Anterolateral margin distinctly con- Etymology. The second author takes
vex, smooth; epibranchial tooth well devel- pleasure in naming the present species after
oped to low, separated from external orbital Mr. Tone Vedenik, the "moving spirit" of
angle by distinct notch; posterolateral mar- the Caving Club "Cmi galeb" in Prebold,
gins strongly converging towards posterior Slovenia, and its expeditions abroad.
carapace margin. Ocular peduncle and cor-
nea slightly reduced; eye occupying about Discussion
% of orbit. Surface of chelipedal carpus
smooth; inner margin with 1 large subla- The caves where crabs had been collect-
melliform tooth and 1 denticle. Ambulatory ed are mostly of modest dimensions. Some
legs relatively short; meri of third and contain sinking streams, while others are
fourth legs about 3.2 and 3.0 times longer primary springs ofpurely hypogean brooks.
than broad respectively; dactylus of third The water temperature on average, was be-
m
and fourth legs subequal to length of prop- tween 22.5°C (at 400 above sea level,
odus. Male abdomen with rectangular seg- below high mountains) and 27-28°C (at sea
ment 6, about 1.1 times longer than broad. level). The pH was generally 7.5-8.0. The
Gl slender, gently curved outwards; termi- food resources in the investigated caves are
nal segment slender, conical, distal part par- very diverse. While some of them are evi-
ticularly slim, about 0.4 times length of dently nutrient-poor, others contain large
subterminal segment. G2 about 1.0 times amounts of plant debris. These are also the
length of Gl; distal segment well devel- ones inhabited by a rich trogloxene fauna.
oped, about 0.5 times length of basal seg- The most frequently observed animals in
ment. — these caves are pigmented or troglomorphic
Discussion. The relatively short ambu- shrimps of the family Atyidae (Caridina
latory legs of S. vedeniki, new species, al- spp.) while Palaemonidae (Macrobrachium
lies it with S. boex, new species. The leg sp.) are generally rare and never troglo-
proportions ofS. vedeniki, however, are still morphic. The relatively large number of
greater than those of S. boex. The Gls of crabs as well as large mollucs (including
S. vedeniki are relatively stouter and the ter- Brotia sp. and Corbicula sp.) found in the
minal segment straighter than those of S. Bohol caves is interesting. It contrasts
boex. The posterolateral margins of S. ve- somewhat with dinaric sinking rivers which
deniki converge towards the posterior car- are usually richer in insect larvae (Sket
apace margin more strongly than those of 1970, 1979).
S. boex, giving it a less squarish appear- With regards to the cavernicolous species
ance. In addition, the carapace of S. veden- discussed here, there are some interesting
iki is distinctly more inflated than that ofS. morphological and ecological trends. Sun-
boex. dathelphusa cavernicola is clearly a com-
Sundathelphusa vedeniki was found in a pletely troglobiomorphic species, with very
cave (Boho sa Bikahan) northwest of An- reduced pigmentation and eyes (Fig. 4g).
tequera which is subjected to periodic re- Sundathelphusa sottoae on the other hand,
surgence of waters. At the end of the rainy is a less troglobiomorphic species, with
season, the water in the cave is stagnant. more well developed eyes (Fig. 4h-l) and
There is a large amount of organic debris- less obvious loss of pigmentation. Sunda-
like tree branches and leaves in the pools, thelphusa urichi has also been found in one
as well as numerous mollusc shells. Live cave only, and while its body is poorly pig-
surface-dwelling gastropods are numerous, mented, the eyes are only slightly reduced
particularly a large species of Brotia. Also (Fig. 4m, p). Sundathelphusa vedeniki has
common is a normally pigmented catfish been found in nutrient rich caves only but
(Clariidae: Clarias sp.). it is probably mainly an epigean species.
702 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OFWASHINGTON
Fig. 2. a-f, Sundathelphusa boex, new species, holotype male (37.4 X 30.9 mm) (NMCR); g, S. sottoae,
new species, paratype female (26.0 X 21.0 mm) (ZRC 1996.1553); h-k, S. sottoae, new species, h-k, holotype
male (17.3 X 13.4 mm) (NSMT-Cr 8938). a, carapace; b, rightthird ambulatory leg; c, rightfourth ambulatory
leg; d, ventral view ofleft Gl; e, dorsal view ofleft Gl; f, left G2; g, carapace; h, right third ambulatory leg;
i, rightfourth ambulatory leg;j, ventral view ofleft Gl; k, left G2 (afterTakeda, 1983) (differentscale fromj).
VOLUME 109, NUMBER 4 703
Fig. 3. a-f,Sundathelphusa urichi, new species, holotypemale (36.6 X 27.9 mm) (NMCR); g-1,S. vedeniki,
new species, holotype male (28.2 X 22.3 mm) (NMCR). a, carapace; b, right third ambulatory leg; c, right
fourth ambulatory leg; d, ventral view ofleftGl; e, dorsal view ofleftGl; f, leftG2; g, carapace; h, rightthird
ambulatory leg; i, right fourth ambulatory leg;j, ventral view ofleft Gl; k, dorsal view ofleft Gl; 1, left G2.
704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON
«;to^
Fig. 4. Orbits and eyes ofSundathelphusa species fromBohol. a-f, 5'. boex: a, holotypemale (37.4 X 30.9mm.
Sierra Bulliones, NMCR); b, male (24.0 X 20.1 mm, Pahangong Talon, ULB); c, paratype male (41.0 X 33.0 mm,
Batuan, ZRC 1996.1549); d,paratypemale (34.2 X 27.8 mm, CastigioCave,ZRC 1996.1550); e,male(34.6 X 28.8
mm, Batuan, NMCR); f, male (10.2 X 8.7 mm, Carmulaon, ULB). g, S. cavemicola: holotypefemale(25.7 by 21.0
mm, Quinapon-anCave, NSMT-Cr8937) (afterTakeda, 1983: Fig. 2). h-1,S. sottoae: h, holotypemale(17.3 X 13.4
mm, Ughob Cave, NSMT-Cr 8938); i, paratype female (26.0 X 21.0mm, Bonugan Cave, ZRC 1996.1553);j, male
(17.4 X 13.8 mm, Batuan, ZRC 1996.1548); k, paratype female (28.0 X 25.0 mm, Bonugan Cave, NMCR); 1, male
(14.8 X 12.5 mm, Kalumpan, ULB); m-p, S. urichi (Quilas Cave): m, holotypemale (36.6 X 27.9 mm, NMCR); n,
paratype male (16.5 X 20.5 nun, ULB); o, paratype male (21.6 X 15.5 mm, ZRC 1996.1555); p, paratype female
(32.6 X 25.0mm,ZRC 1996.1556).q,r,S. vedeniki(BohosaBikahan): q,holotypemale(28.2 X 22.3mm,NMCR);
r, paratype male (33.9 X 26.5 mm, ZRC 1996.1552).
