THE RAFFLES BULLETIN OF ZOOLOGY 2009
THE RAFFLES BULLETIN OF ZOOLOGY 2009 57(1): 65–89
Date of Publication: 28 Feb.2009© National University of Singapore

EPIGEAN AND HYPOGEAN FRESHWATER SHRIMPS OF BOHOL ISLAND,CENTRAL PHILIPPINES (CRUSTACEA: DECAPODA: CARIDEA)
Yixiong Cai
National Biodiversity Centre, National Parks Board, 1 Cluny Road, Singapore 259569, Republic of Singapore
Email: caiyixiong@yahoo.com (Corresponding author)

Satish Choy
Queensland Department of Natural Resources and Water, 120 Meiers Road, Indooroopilly, Queensland 4068, Australia
Email: choys@nr.qld.gov.au

Peter K. L. Ng
Tropical Marine Science Institute, National University of Singapore, Republic of Singapore
Email: dbsngkl@nus.edu.sg

ABSTRACT. – The epigean and hypogean freshwater shrimps from Bohol Island, central Philippines, collectedduring various speleological expeditions by the Philippine, Slovenian and Singaporean scientists between
1995 and 2000 are reported here. The material contains 25 species of caridean shrimps in two families.
The Atyidae are represented by 16 species from four genera: Antecaridina, Halocaridinides and Atyopsis,
each with one species; and Caridina, with 13 species, including seven new taxa. The Palaemonidae arerepresented by two genera, Macrobrachium with eight species and Palaemon with one. Detailed descriptions
are provided for new and rarely reported species.
KEY WORDS. – Bohol, Philippine, freshwater shrimp, Atyidae, Palaemonidae, new species.

INTRODUCTION

Biodiversity Research, the National University of Singapore,and the Department of Biology, University of San Carlos,
Cebu, in December 2000, resulted in the identification of 25
species of caridean shrimps: 16 species of Atyidae, including
seven new species and nine species of Palaemonidae. In thisstudy, detailed descriptions and illustrations are provided
for the new and the rarely reported species, and brief notes
are provided for the more common and widely distributed
species.

The freshwater shrimps of the Philippine Archipelago havebeen comparatively well studied (Dana, 1852; Cowles, 1915a,
1915b; Blanco, 1935, 1939; Holthuis, 1950; 1978; Johnson,
1962; Chace, 1983; 1997; Balete & Holthuis, 1992; Chace& Bruce, 1993; Sket, 1997; Cai & Anker, 2004; Cai &Shokita, 2006a). Prior to this study, 52 species of freshwater
caridean shrimps were reported from the Philippines. The
family Atyidae was represented by seven genera with 28
species: Antecaridina, Halocaridinides, Atyoida, Atyopsis,
Parisia, Edoneus, with one species each, and Caridina, with22 species. The family Palaemonidae was represented by
two genera with 22 species: Macrobrachium with 20 species
and Palaemon with two species. The family Alpheidae wasrepresented by two species of Potamalpheops.

All the specimens discussed in this study were collectedfrom various localities on Bohol Island, the Philippines (Fig.
1). Specimens examined are deposited in the Zoological
Reference Collection, the Raffles Museum of Biodiversity
Research, the National University of Singapore, Singapore(ZRC); Department of Biology, University of San Carlos,
Cebu, Philippines (USC); and Department of Biotechnology,
University of Ljubljana, Slovenia (UOL). The abbreviation cl
is used for carapace length measured in mm from the postorbital margin to the posterior margin of the carapace.

The examination of the recent collection from the Bohol
Island (Fig. 1), central Philippines, collected by Dr. Boris
Sket and his Slovenian colleagues in February 1995 andMarch 1999, and a joint expedition by the Raffles Museum of

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SYSTEMATIC ACCOUNT

Atyopsis Chace, 1983

Atyidae De Haan, 1849

Atyopsis spinipes (Newport, 1847)

Antecaridina Edmondson, 1954

Atya spinipes Newport, 1847: 159 [type locality: Philipine
Islands]
Atya moluccensis – De Man, 1902: 893 (not Atya moluccensis De
Haan, 1849).
Atyopsis spinipes – Chace, 1983: 35, Figs. 20–22; 1997: 4; Cai &Shokita, 2006b: 2133.

Antecaridina lauensis (Edmondson, 1935)
Mesocaris lauensis Edmondson, 1935a: 13, Fig. 4 [type locality:
Namuka and Wangava Island, Lau Archipelago, Fiji];
Edmondson, 1935b: 4.
Antecaridina lauensis – Edmondson, 1954: 368; Holthuis, 1955:
25, Fig. 8d–f; 1956: 51; 1963: 267; 1965: 4, Figs. 2a–r; 1973:
19; Shokita, 1975: 118; 1979: 201; Suzuki, 1980: 47, Figs.
1–5; Smith & Williams, 1981: 49; Hayashi, 1989a: 378, Figs.
172–174; Sket, 1997: 62; Leberer & Cai, 2003: 355; Cai &Shokita, 2006b: 2124.

Material examined. – Two females, cl 6.0–7.2 mm, ZRC 2007.0268,
River outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16
Dec.2000.

Remarks. – Chace (1997) reported this species from Luzon,
Catanduanes, Cebu and a location that probably Mindoro.
De Man (1902) reported Atya moluccensis from Halmahera,for which, Chace (1983) commentd that De Man’s (1902)
specimens “probably represent Atyopsis spinipes, as indicatedby the fewer (3–5) ventral teeth.” According to Chace(1983), the most reliable characters used to separate Atyopsis
moluccensis from A. spinies is the large number of ventral
rostral teeth as 7–16 (vs. 2–6). Atyopsis spinipes occurs
from the Philippines and eastern Lesser Sunda Islands (atca.120°00'E) northwards to Taiwan and Tokuno-shima in the
Ryukyus, and eastwards as far as Samoa (Chace, 1983).

Material examined. – One male, cl 2.8 mm, 16 females, cl 1.8–2.6
mm, 12 juveniles, ZRC 2007.0267, Cave Hinaganan, Panglao, coll.
Y. Cai et al., 17 Dec.2000; 60 juveniles, UOL, Cave Hinaganan,
Panglao, coll. B. Sket, Feb.1995.

Remarks. – Antecaridina lauensis, an anchialine species
with a wide but disjunct distribution, has been reported
from the Lau Archipelago, Fiji Islands, (Edmondson,
1935a, b), Europa Island, Madagascar (Holthuis, 1965),
Sinai Peninsular, Red Sea (Holthuis, 1963), Hawaii Island(Holthuis, 1973), Solomon Islands (Smith & Williams, 1981),
Guam (Leberer & Cai, 2003) and the Ryukyus (Shokita,
1975; 1979; Suzuki, 1980). It has previously been reported
from Bohol by Sket (1997).

Caridina H. Milne Edwards, 1837
Caridina typus H. Milne Edwards, 1837
Caridina typus H. Milne Edwards, 1837: 363 [type locality:
unknown]; Holthuis, 1965: 10, Fig. 3; Chace, 1997: 21; Cai &Anker, 2004: 236; Wowor et al., 2004: 341; Fig. 5F; Cai et al.,
2006: 412, Figs. 13–15; Cai & Shokita, 2006b: 2134.

Halocaridinides Fujino & Shokita, 1975
Halocaridinides trigonophthalma(Fujino & Shokita, 1975)

Material examined. – One female, cl 3.2 mm, ZRC 2007.0269, river
outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000;
1 male, cl 4.7 mm, 8 females, cl 2.8–4.0 mm, 2 ovigerous females,
cl 6.8–7.2 mm, ZRC 2007.0270, River outside Inabacan Cave,
Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, cl 5.7 mm, 3
ovigerous females, cl 6.6–7.8 mm, ZRC 2007.0271, Loboc river,
Loboc, coll. Y. Cai et al, 19 Dec.2000.

Halocaidina (Halocaridinides) trigonophthama Fujino & Shokita,
1975: 106, Figs. 7, 8 [type locality: a well in Okinawa Island,
Ryukyus, Japan]; Shokita, 1979: 201.
Palauatya dasyomma Hart, 1980: 481, Figs. 1–31 [type locality:
Anguar Island, Palau, Caroline Islands].
Halocaridinides trigonophthalma – Holthuis, 1982: 31, Figs. 3a–u;
Gurney, 1984: 591; Hayashi, 1989b: 497, Fig. 177; Shokita,
2003: 249, Fig. 17C; Naruse et al., 2003: 1; Cai & Shokita,
2006b: 2132.

Remarks. – Caridina typus is a very common insular or
coastal freshwater species, which is widely distributed in
the islands of Indo-West Pacific region.

Material examined. – Eight juveniles, incomplete, UOL, Alijauan
Bridge, stream, S. Duero, coll. B. Sket, 5 Mar.1999.

Caridina villadolidi Blanco, 1939

Remarks. – The taxonomy of Halocaridinides trogonophthalma
has recently been discussed by Naruse et al. (2003) and Cai& Shokita (2006b). The present record is based on eight
incomplete juvenile specimens, all of which were collectedfrom an interstitial habitat. These specimens, however, clearlyshow the characteristic of the species, i.e. the extremely shortrostrum, the slender pereiopods, the fewer uropodal teeth
as well as the short and stout telson. This represents a newrecord for the Philippines.

Caridina typus var. longirostris De Man, 1892: 370, Pl. 22: 22f–i[type locality: Palopo, Sulawesi (Celebes), Indonesia].
Caridina villadolidi Blanco, 1939: 389, Pl. 1 [type locality: Laoag
River, Luzon, the Philippines] Hung et al., 1993: 485, Fig. 3;
Chace, 1997: 21, Fig. 12; Cai & Ng, 2001: 668, Fig. 4; Cai& Shokita, 2006a: 248.
Caridina typus – De Silva, 1982: 135. Figs. 4a–h (not Caridina
typus H. Milne Edwards, 1837).

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THE RAFFLES BULLETIN OF ZOOLOGY 2009Material examined. – Ten females, cl 3.3–3.8 mm, ZRC 2007.0272,
river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16
Dec.2000; 1 male, cl 3.4 mm, 11 females, cl 3.4–4.4 mm, ZRC
2007.0273, river outside Inabacan Cave, Antequrra, coll. Y. Cai
et al., 18 Dec.2000; 1 male, cl 4.9 mm, ZRC 2007.0274, Loboc
River, Loboc, coll. Y. Cai et al., 19 Dec.2000.

mm, ZRC 2007.0276, river outside Inabacan Cave, Antequrra, coll.
Y. Cai et al., 18 Dec.2000.

Remarks. – Chace (1997) commented that Caridina
laoagensis will eventually fall into synonymy with thevariable C. weberi De Man, 1892. Cai & Anker (2004) has
shown that C. laoagensis differs markedly from all subspeciesof C. weberi and other congeners, and should be regarded
as a good species. Caridina laoagensis is characterized byits straight rostrum which is slightly crested basally above
the orbit, and the pattern of distal spines and setae on thetelson, on which, the median plumose seatae are inserted
betweem two lateral spines. This is one of the most common
species of atyids in the Philippines.

Remarks. – Caridina villadolidi has been reported from
Taiwan (Hung et al., 1993), the Philippines (Blanco, 1939;
Chace, 1997; Cai & Shokita, 2006a), Sulawesi and Halmahera
in Indonesia (Cai & Ng, 2001), and Sri Lanka (De Silva,
1982, misidentified as Caridina typus).

Caridina laoagensis Blanco, 1939
Caridina laoagensis Blanco, 1939: 390, Pl. 2 [type locality: Laoag
River, Province of Ilocos Norte, Luzon, the Philippines]; Chace,
1997: 12; Cai & Anker, 2004: 237, Fig. 3; Cai & Shokita,
2006a: 248; Cai & Shokita, 2006b: 2141.

Caridina endehensis De Man, 1892
Caridina brevicarpalis var. endehensis De Man, 1892: 399, Pl. 24
Fig. 30e [type locality: Nuawari, near Ende, Flores, Indonesia];
Bouvier, 1925: 34; Roux, 1928: 218; Blanco, 1935: 34, Pl. 2
Fig. 25; Chace, 1997: 8, Fig. 3.
Caridina brevicarpalis – Holthuis, 1978: 38 (not Caridina
brevicarpalis De Man, 1892).

Material examined . – Two males, cl 2.6–2.9 mm, 1 female, cl
2.9 mm, 2 ovigerous females, cl 5.6–6.0 mm, ZRC 2007.0275,
river outside the Inabacan Cave, Antequrra, coll. Y. Cai et al., 16
Dec.2000; 3 males, cl 3.2–3.7 mm, 2 ovigerous females, cl 6.2–6.3

Fig. 1. Map of the Bohol Island, Philippines, showing the collection sites.

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Cai et al.: Freshwater shrimps from Bohol
Caridina endehensis – Wowor et al., 2004: 341, Fig. 5M; Cai &Shokita, 2006a: 248.

Remarks. – Cai & Ng (2007) recently reviewed the taxonomyof the Caridina gracilirostris species group in detail, referring
the form with no appendix interna in endopod of male firstpleopod as C. gracilirostris sensu stricto and the other,with a distinct appendix interna in the endopod of malefirst pleopod, as C. appendiculata Jalihal & Shenoy, 1998.
The specimens from Bohol have no appendix interna on theendopod of the male first pleopod, and are here referred to
C. gracilirostris. It is an insular or coastal freshwater species
commonly distributed in Indo-West Pacific region.

Material examined. – 13 males, cl 3.0–4.2 mm, 25 females, cl
2.7–6.2 mm, 2 ovigerous, females, cl 4.6–6.2 mm, ZRC 2007.0277,
river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16
Dec.2000; 11 males, cl 3.2–4.6 mm, 13 females, cl 2.9–4.8 mm, 2
ovigerous females, cl 5.3–6.4 mm, ZRC 2007.0278, river outside
Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000.

Remarks. – Holthuis (1978) reported Caridina brevicarpalis
from Sumba, Lesser Sunda Islands, Indonesia. According to
his description, his specimens “can unhesitatingly be brought
to ssp. endehensis”. Caridina endehensis has previously
been reported from the Philippines by Blanco (1935),
Chace (1997) and Cai & Shokita (2006a). It is distributed
from Philippines to Flores and Sumba in Indonesia. The
prominently elongated rostrum easily separates C. endehensis
from C. brevicarpalis and does not appear to be variable.
We therefore follow Cai & Shokita (2006a) and regard thetaxon as a full species.

Caridina lobocensis, new species(Figs. 2–4)
Material examined. – Holotype: male, cl 5.7 mm, USC, tributaryof Loboc River, Loboc, coll. Y. Cai, 19 Dec.2000.
Paratypes: Seventy males, cl 3.8–5.3 mm, 19 females, cl 3.9–7.5
mm, 83 ovigerous females, cl 5.3–7.2 mm, ZRC 2007.0283, data
same as holotype.

Description. – Rostrum (Figs. 2A, 3A) straight, reachingslightly beyond end of antennular peduncle; rostral formula:
2–4+10–12/3–7. Antennal spine fused with inferior orbital
angle. Pterygostomian margin subrectangular.

Caridina elongapoda Liang & Yan, 1977
Caridina nilotica elongapoda Liang & Yan, 1977: 220, Figs. 5–8[type locality: Xinzai, Gulei village, Zhangpu County, Fujian,southern China].
Caridina aff. brachydactyla - Yeo et al., 1999: 218, Figs. 10–14;
Wowor et al., 2004: Fig. 6B;
Caridina elongapoda Cai & Shokita, 2006a: 249.

Sixth abdominal somite 0.44 times of carapace, 1.4 times
as long as fifth somite, distinctly shorter than telson. Telson(Figs. 3B, 4A) 2.6 times as long as wide, terminating in a
projection, with 3 or 4 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of spines distinctly shorter
than intermediate pairs of setae. Preanal carina (3I) high,lacking spine.

Material examined. – Two males, cl 4.1–4.5 mm, 3 ovigerous
females, cl 5.6–5.8 mm, ZRC 2007.0279, tributary of Loboc River,
Loboc, coll. Y. Cai, 19 Dec.2000; 1 male, cl 4.0 mm, 5 females, cl
2.8–3.4 mm, 3 ovigerous females, cl 5.3–6.4 mm, ZRC 2007.0280,
river outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 18
Dec.2000; 6 males, cl 3.1–4.7 mm, 17 females, cl 2.8–4.3 mm,
ZRC 2007.0281, river outside Inabacan Cave, Antequrra, coll. Y.
Cai et al., 16 Dec.2000.

Eyes developed, anterior end reaching to 0.8 times length ofbasal segment of antennular peduncle. Antennular peduncle
0.55 times as long as carapace; basal segment of antennularpeduncle as long as both of second and third segment
lengths, anterolateral angle reaching 0.20 times length ofsecond segment, second segment distinctly longer than third
segment. Stylocerite reaching slightly beyond end of basal
segment of antennular peduncle. Scaphocerite (Fig. 2B) 2.5
times as long as wide.

Remarks. – Caridina elongapoda has recently been reported
from the Philippines (Cai & Shokita, 2006a). Cai & Shokita(2006a) reassigned the material identified as Caridina aff.
brachydactyla by Yeo et al. (1999) from Pulau Tioman,
Malaysia, to this species.Current known distribution issouthern China, Malaysia and the Philippines.

Incisor process of mandible (Fig. 2C) ending in irregularteeth, molar process truncated. Lower lacinia of maxillula(Fig. 2D) broadly rounded, upper lacinia elongated, witha number of distinct teeth on inner margin, palp slender.
Upper endites of maxilla (Fig. 2E) subdivided, palp short,scaphognathite tapering posteriorly with some long, curved
setae at posterior end. Palp of first maxilliped (Fig. 2F)
broadly triangular. Second maxilliped (Fig. 2G) typicalof genus. Third maxilliped (Fig. 2H) reaching to end ofantennular peduncle, with ultimate segment shorter thanpenultimate segment.

Caridina gracilirostris De Man, 1892
Caridina gracilirostris De Man, 1892: 399, Pl. 25 Fig. 31–31d[type locality: Balangnipa, Sulawesi (Celebes) Indonesia];
Bouvier, 1925: 142, Figs. 305–307; Holthuis, 1965: 23, Fig.
7; Tiwari & Pillai, 1971: 83, Fig. 2a, b; Chace, 1997: 10, Fig.
4; Wowor et al., 2004, Fig. 5Q; Cai & Shokita, 2006a: 250;
Cai & Shokita, 2006b: 2135; Cai & Ng, 2007: 1586.
Material examined. – Two males, cl 2.5–2.6 mm, 3 females, cl
2.0–3.3 mm, 1 ovigerous female, cl 3.7 mm, ZRC 2007.0282,
tributary of Loboc River, Loboc, coll. Y. Cai, 19 Dec.2000.

Epipods on first 4 pereiopods. First pereiopod (Figs. 3C, 4D)
reaching to end of basal segment of antennular peduncle;
merus 1.7–2.1 times as long as broad, slightly longer than68

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Fig. 2. Caridina lobocensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, scaphocerite; C, mandible; D,maxillula; E, maxilla; F, first maxilliped; G, second maxilliped; H, third maxilliped. Scale bars: A = 2 mm, B–H = 1 mm (female, cl 6.9mm, Paratype, ZRC, Philippines).

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Cai et al.: Freshwater shrimps from Bohol

Fig. 3. Caridina lobocensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, first
pereiopod; D, second pereiopod; E, third pereiopod; F, dactylus of third pereiopod; G, fifth pereiopod; H, dactylus of fifth pereiopod;
I, preanal carina; J, uropodal diaeresis. Scale bars: A = 2 mm, B, F, H, J = 0.2 mm; C–E, G = 1 mm; I = 0.5 mm (male, cl 5.7 mm,paratype, ZRC, Bohol Island, Philippines).

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Fig. 4. Caridina lobocensis, new species: A, telson; B, endopod of male first pleopod; C, appendix masculina and appendix interna ofmale second pleopod; D, first pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fifth pereiopod; I,
dactylus of fifth pereiopod. Scale bars: A, D–F, H = 1 mm; B, C, G, I = 0.2 mm (A–C, male, cl 5.7 mm, paratype, ZRC, Bohol Island,
Philippines; D–I, female, cl 6.9 mm, paratype, ZRC, Bohol Island, Philippines).

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Cai et al.: Freshwater shrimps from Bohol
carpus; carpus excavated anteriorly, shorter than chela, 1.1–
1.2 times as long as high; chela 2.2 times as long as broad;fingers distinctly shorter than palm. Second pereiopod (Figs.
3D, 4E) reaching to end of second segment of antennularpeduncle; merus as long as carpus, 4.2–5.0 times as long as
broad; carpus 1.1 times as long as chela, 2.3–2.4 times aslong as high; chela 2.3–2.4 times as long as broad; fingers 1.6
times as long as palm. Third pereiopod (Figs. 3E, F, 4F, G)
reaching slightly beyond end of scaphocerite, propodus 9–11
times as long as broad, 5.2–5.7 times as long as dactylus;
dactylus 2.7–3.0 times as long as wide (spines included),terminating in 2 very strong claws, with 4–6 accessory
spines on its flexor margin. Fifth pereiopod (Figs. 3G, H,
4H, I) reaching beyond end of second segment of antennularpeduncle, propodus 13–18 times as long as broad, 5.5–5.6
times as long as dactylus, dactylus 2.7–3.0 times as long as
wide (spinules included), terminating in 2 large claws, with
26–30 spinules on its flexor margin.

of the antennular peduncle vs. does not reaches) (Figs. 2,
3; cf. Cai & Shokita, 2006b: Figs. 13, 14).

Caridina liaoi, new species(Figs. 5, 6)
Material examined. – Holotype: Ovigerous female, cl 4.1 mm,
USC, Bilar River, Bilar, coll. Y. Cai et al., 19 Dec.2000.
Paratypes: 82 males, cl 2.8–4.3 mm, 112 females, cl 3.7–5.0 mm,
26 ovigerous females, cl 4.2–5.0 mm, ZRC 2007.0284, data same
as holotype.

Description. – Rostrum (Figs. 5A, 6A) long, reaching nearto or slightly beyond end of scaphocerite; rostral formula:
3–4+13–14/13–15. Antennal spine fused with inferior orbital
angle. Pterygostomian margin broadly rounded.
Sixth abdominal somite 0.51 times of carapace, 1.7 times
as long as fifth somite, distinctly shorter than telson. Telson(Figs. 6B, C) 3.5 times as long as wide, not terminating in a
projection, with 3 or 4 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of spines distinctly longer
than intermediate pairs of spiniform setae. Preanal carina(Fig. 6K) high, triangular, lacking spine.

Endopod of male first pleopod (Fig. 4B) subtriangular, half
length of exopod, appendix interna stout, exceeding end of
endopod. Appendix masculina of male second pleopod (Fig.
4C) 2/3 length of endopod, with appendix interna reachingbase of distal one-third of appendix masculina.
Uropodal diaeresis (Fig. 3J) with 18 movable spinules.

Eyes well developed, anterior end reaching to 0.8 times
length of basal segment of antennular peduncle. Antennularpeduncle 0.74 times as long as carapace; basal segmentof antennular peduncle longer than both second and third
segment lengths, anterolateral angle reaching 0.20 times
length of second segment, second segment distinctly longer
than third segment. Stylocerite reaching to 0.8 times lengthof basal segment of antennular peduncle. Scaphocerite (Fig.
6D) 3.4 times as long as wide.

Ovigerous females with eggs sized 0.45 × 0.25 mm.
Habitat. – Caridina lobocensis, new species, was collectedfrom a tributary of the Loboc River near Loboc town, Bohol
Island in central Philippines. The shrimps were living among
submerged grass at edges of the tributary.
Etymology. – The new species is named after its type localityLoboc River, Bohol Island, the Philippines.

Incisor process of mandible ending in irregular teeth, molar
process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongated, with a number of distinct
teeth on inner margin, palp slender. Upper endites of maxilla
subdivided, palp short, scaphognathite tapering posteriorly
with some long, curved setae at posterior end. Palp of first
maxilliped broadly triangular. Second maxilliped typicalof genus. Third maxilliped reaching to end of antennularpeduncle, with ultimate segment as long as penultimate
segment.

Remarks. – Based on the form of the rostrum, the highpreanal carina, the large number of uropodal teeth and theform of the telson, C. lobocensis, new species, belongs to
the C. weberi species group. In the group, it is most similar
to C. okinawa Cai & Shokita, 2006b, from Okinawa Islandof the Ryukyu Islands, southern Japan, in term of the rostralformula, the long stylocerite, the broad scaphocerite, the stout
first pereiopod, and its short fingers, and the third to fifth
pereiopods ending in two claws. However, it could be easily
separated from C. okinawa by the well developed eyes (vs.
eyes less developed, with a short eye stalk and small cornea)
and the longer rostrum which reaches slightly beyond endof antennular peduncle (vs. reach to end of second segmentof antennular peduncle) (Figs. 2–4; cf. Cai & Shokita,
2006b: Fig. 12). Caridina lobocensis also resembles another
Ryukyuan species, C. macrodentata Cai & Shokita, 2006b,in the form of the dactylus of the third to fifth pereiopods,
Nevertheless, it could be differentiated from the latter by
the longer rostrum, which reaches slightly beyond the endof the antennular peduncle (vs. reaches middle of the second
segment of the antennular peduncle) and the long stylocerite(reaches to or slightly beyond the end of the basal segment

Epipods on first 4 pereiopods. First pereiopod (Figs. 5B, 6E)
reaching to end of basal segment of antennular peduncle;
merus 1.9–2.2 times as long as broad, shorter than carpus;
carpus excavated anteriorly, shorter than chela, 1.3–1.7 times
as long as high; chela 2.3–3.5 time as long as broad; fingers
slightly shorter, as long as or distinctly longer than palm.
Second pereiopod (Figs. 5C, 6F) reaching to end of second
segment of antennular peduncle; merus as long as carpus,
4.2–5.0 times as long as broad; carpus 1.1 times as long aschela, 2.3–2.4 times as long as high; chela 2.3–2.4 times
as long as broad; fingers 1.6 times as long as palm. Third
pereiopod (Figs. 5D, E, 6G, H) reaching to end of antennular

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Fig. 5. Caridina liaoi, new species. A: cephalothorax and cephalic appendages, lateral view; B, first pereiopod; C, second pereiopod; D,
third pereiopod; E, dactylus of third pereiopod; F, fifth pereiopod; G, dactylus of fifth pereiopod; H, endopod of male first pleopod; I,
appendix masculina and appendix interna of male second pleopod. Scale bars: A–D, F = 1 mm; E, G = 0.1 mm; H, I = 0.2 mm (male,cl 4.0 mm, paratype, ZRC, Bilar river).

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Fig. 6. Caridina liaoi, new species, A: cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, telson; D,
scaphocerite; E, first pereiopod; F, second pereiopod; G, third pereiopod; H, dactylus of third pereiopod; I, fifth pereiopod; J, dactylus of
fifth pereiopod; K, preanal carina; L, uropodal diaeresis. Scales: A, C–G, I = 1 mm; B, L = 0.2 mm; H, J = 0.1 mm; K = 0.5 mm. (ovig.
females, cl 4.6 mm, paratype, ZRC, Bilar River).

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Antequrra, coll. Y. Cai et al., 16 Dec.2000; 41 males, cl 2.6–3.5
mm, 38 females, cl 2.9–4.1 mm, ZRC 2007.0287, Cave Bongkawi,
Canla-As village, Antequrra, coll. Y. Cai.et al., 16 Dec.2000; 3
males, cl 3.0–3.8 mm, 11 females, cl 2.3–3.8 mm, ZRC 2007.0288,
spring outside cave Bones, Canla-As village, Antequrra, coll. P.
K. L. Ng & L. Liao, 16 Dec.2000; 6 males, cl 2.8–3.4 mm, ZRC
2007.0289, spring outside Cave Inabacan, Antequrra, coll. Y.
Cai et al., 18 Dec.2000; 22 males, cl 3.0–4.0 mm, 37 females,
cl 2.2–3.3 mm, 15 juveniles, ZRC 2007.0290, Cave Serpia,
Ginguyuran, Dimiao, coll. B. Sket, Feb.1995; 1 juvenile, ZRC
2007.0291, Cave Camulaon, lower part, Cugon, Jagna, coll. B.
Sket, Feb.1995; 7 juveniles, ZRC2007.0292 , spring no.1, Roxas
Park, Garcia-Hernandez, coll. B. Sket, Feb.1995; 4 juveniles,
ZRC 2007.0293, spring no. 2, Roxas Park, Garcia-Hernandez,
coll. B. Sket, Feb.1995; 6 juveniles, ZRC 2007.0294, spring
Anislaq, outside, Valencia, coll. B. Sket, Feb.1995; 2 juveniles,
ZRC 2007.0295, spring below Cave Serpia, Ginguyuran, Dimiao,
B. Sket, Feb.1995; 23 juveniles, ZRC 2007.0296, spring Anislaq,
outside, Velencia, coll. B. Sket, Feb.1995.

peduncle, propodus 11–12 times as long as broad, 4.5–4.7
times as long as dactylus; dactylus 3.1–3.3 times as long as
wide (spines included), with 4–6 accessory spines on flexor
margin. Fifth pereiopod reaching to end of second segmentof antennular peduncle, propodus 14–18 times as long as
broad, 3.6–3.8 times as long as dactylus, dactylus 4.0–4.4
times as long as wide (spinules included), terminating in 1
large claw, with 60–65 spinules on flexor margin.
Endopod of male first pleopod (Fig. 5H) sub-rectangular,
two-fifth length of exopod, no appendix interna. Appendix
masculina of male second pleopod (Fig. 5I) half lengthof endopod, with appendix interna reaching near end of
appendix masculina.
Uropodal diaeresis (Fig. 6L) with 14–16 movablespinules.
Ovigerous females with eggs sized 0.92–0.95 × 0.55–0.60
mm.

Description. – Rostrum (Figs. 7A, 8A) very short, reachingnear end of basal segment of antennular peduncle; unarmed
dorsally or armed dorsally with indistinct small teeth, unarmed
ventrally. Antennal spine indistinct, fused with inferior orbital
angle. Pterygostomian margin broadly rounded.

Habitat. – Caridina liaoi, new species, was collected froma tributary of the Bilar River near Bilar town, Bohol Islandin central Philippines.

Sixth abdominal somite 0.45 times of carapace, 1.8 times
as long as fifth somite, as long as telson. Telson (Figs.
7B, C) 2.8 times as long as wide, not terminating in a
projection, with 3 or 4 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of spines distinctly longer
than intermediate pairs. Preanal carina high, sub-triangular,
lacking spine.

Etymology. – The new species is named after Dr. Lawrence
Liao, who has been instrumental in helping the first author
making collections in the Philippines for the present
study.
Remarks.- Caridina liaoi, new species, is similar to C.
buhi Cai & Shokita, 2006a, from Luzon, by the form of
pereiopods, but it differs from C. buhi by its longer rostrum(reaches to or slightly beyond the anterior margin of thescaphocerite vs. reaches to the end of the second segmentof the antennular peduncle or to the end of the antennularpeduncle); the more ventral rostral teeth (13–15 vs. 3–7); thetelson does not terminate in a projection vs. terminate in a
projection; the bigger appendix interna on the male second
pleopod (reaches near the end of the appendix masculina vs.
reaches the base of the distal two-third length of the appendix
masculina) and the distal spines on the telson (lateral spines
much longer than the intermediate pairs of spiniform setae
vs. lateral spines subequal to intermediate pairs) (Figs. 5,
6; cf. Cai & Shokita, 2006a: Figs. 3, 4).

Eyes reduced, anterior end reaching to 0.3–0.4 times lengthof basal segment of antennular peduncle. Antennularpeduncle 0.64 times as long as carapace; basal segmentof antennular peduncle shorter than both second and third
segment lengths, anterolateral angle reaching 0.20 times
length of second segment, second segment distinctly longer
than third segment. Stylocerite reaching to 0.8 times lengthof basal segment of antennular peduncle. Scaphocerite (Fig.
7D) 3.5 times as long as wide.
Incisor process of mandible (Fig. 7E) ending in irregularteeth, molar process truncated. Lower lacinia of maxillula(Fig. 7F) broadly rounded, upper lacinia elongated, witha number of distinct teeth on inner margin, palp slender.
Upper endites of maxilla (Fig. 7G) subdivided, palp short,scaphognathite tapering posteriorly with some long, curved
setae at posterior end. Palp of first maxilliped (Fig. 7H)
broadly triangular. Second maxilliped (Fig. 7I) typicalof genus. Third maxilliped (Fig. 7J) reaching to end ofantennular peduncle, with ultimate segment slightly shorter
than penultimate segment.

Caridina boholensis, new species(Figs. 7, 8)
Material examined. – Holotype: Male cl 3.9 mm, USC, Quila Cave,
Nueva Vida Norte, Batuan, coll. B. Sket, Feb.1995.
Paratypes: 87 males, cl 3.0–4.2 mm, 115 females, cl 2.7–4.7 mm,
1 ovigerous females, 3.8 mm, eggs 0.75 x 0.50 mm, UOL, data
same as holotype; 10 males, cl 3.0–4.2 mm, 10 females, cl 2.7–4.7mm, 1 ovigerous females, 4.6 mm, eggs 0.75 × 0.50 mm, ZRC
2007.0285, data same as holotype.

Epipods on first four pereiopods. First pereiopod (Figs. 7K,
8B) reaching beyond end of basal segment of antennularpeduncle; merus 2.3 times as long as broad, shorter thancarpus; carpus excavated anteriorly, shorter than chela, 2.0
times as long as high; chela 2.3 times as long as broad;fingers slightly longer than palm. Second pereiopod (Figs.

Others: Nine males, cl 2.8–3.8 mm, 5 females, cl 37–3.8 mm,
11 juveniles, ZRC 2007.0286, Cave Bones, Canla-As village,75

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Fig. 7. Caridina boholensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson;
D, scaphocerite; E, mandible; F, maxillula; G, maxilla; H, first maxilliped; I, second maxilliped; J, third maxilliped; K, first pereiopod;
L, second pereiopod. Scale bars: A, B, D = 1 mm, E–L = 0.5 mm, C = 0.1 mm (male, cl 3.7 mm, paratype, ZRC, Cave Bongkawi,
Antequrra).

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Fig. 8. Caridina boholensis, new species: A, cephalothorax and cephalic appendages, lateral view; B, first pereiopod; C, second pereiopod;
D, third pereiopod; E, dactylus of third pereiopod; F, fifth pereiopod; G, dactylus of fifth pereiopod; H, male first pleopod; I, appendix
masculina and appendix interna of male second pleopod; J, uropodal diaeresis. Scale bars: A = 1 mm; B–D, F = 0.5 mm; E, G = 0.1 mm;
H–J = 0.2 mm (A–C, ovig. females, cl 3.8 mm, paratype, ZRC; D–J, male, cl 3.7 mm, paratype, ZRC, Cave Bongkawi, Antequrra).

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basal segment of antennular pedunlcle; unarmed. Antennal
spine fused with inferior orbital angle. Pterygostomian
margin sub-rectangular.

7L, 8C) reaching to end of scaphocerite; merus distinctly
shorter than carpus, 4.4 times as long as broad; carpus 1.2
times as long as chela, 6.9 times as long as high; chela 3.7
times as long as broad; fingers 1.6 times as long as palm.
Third pereiopod (Figs. 8D, E, 9E, F) reaching beyond endof scaphocerite by length of dactylus, propodus 10 times aslong as broad, 3.7 times as long as dactylus; dactylus 3.0
times as long as wide (spines included), with 4 or 5 accessory
spines on flexor margin. Fifth pereiopod (Figs. 8F, G, 9G,
H) reaching to end of scaphocerite, propodus 15 times aslong as broad, 3.6 times as long as dactylus, dactylus 3.4
times as long as wide (spinules included), terminating in 1
large claw, with 62 spinules on flexor margin.

Sixth abdominal somite 0.43 times of carapace, 1.8 times
as long as fifth somite, slightly shorter than telson. Telson(Fig. 9B) 2.7 times as long as wide, not terminating in a
projection, with 4 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of distal spines distinctly
longer than intermediate pairs. Preanal carina low, subtriangular, lacking spine.
Eyes well developed, anterior end reaching to 0.7 times
length of basal segment of antennular peduncle. Antennularpeduncle 0.63 times as long as carapace; basal segment ofantennular peduncle distinctly longer than both second andthird segment lengths, anterolateral angle reaching 0.20 times
length of second segment, second segment distinctly longer
than third segment. Stylocerite reaching to 0.7 times lengthof basal segment of antennular peduncle. Scaphocerite 3.6
times as long as wide.

Endopod of male first pleopod (Fig. 8H) sub-triangular, aquarter length of exopod, no appendix interna. Appendix
masculina of male second pleopod (Fig. 8I) half length ofendopod, with appendix interna 0.4 times length of appendix
masculina.
Uropodal diaeresis (Fig. 8J) with 17 movable spinules.

Incisor process of mandible ending in irregular teeth, molar
process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongated, with a number of distinct
teeth on inner margin, palp slender. Upper endites of maxilla
subdivided, palp short, scaphognathite tapering posteriorly
with some long, curved setae at posterior end. Palp of first
maxilliped rounded, exopod with short flagellum. Endopodof second maxilliped slender. Third maxilliped reaching
beyond end of scaphocerite, with ultimate segment slightly
longer than penultimate segment.

Ovigerous females with eggs sized 0.75–0.85 × 0.50–0.60mm.
Habitat. – Caridina boholensis, new species, was collectedfrom several locations of subterranean water in Bohol, either
from cave streams or spring water near caves.
Etymology. – The new species is named after its type
locality-Bohol Island, the Philippines.
Remarks. – Caridina boholensis, new species, is most similar
to C. cebuensis Cai & Shokita, 2006, from Cebu Island,
central Philippines. However, it could be differentiated by
the much reduced eyes, the slender carpus of first pereiopods(2.0 times as long as high vs. 1.6 times in C. cebuensis) andthe slender carpus of second pereiopod (6.9 times as long as
high vs. 4.4 times). Caridina boholensis is the most common
subterreanean Caridina species in Bohol Island.

Epipods on first 4 pereiopods. First pereiopod (Fig. 9C)
stout, reaching to end of second segment of antennularpeduncle; merus 1.6 times as long as broad, slightly shorter
than carpus; carpus excavated anteriorly, shorter than chela,
1.3 times as long as high; chela 1.6 times as long as broad;fingers slightly shorter than palm. Second pereiopod (Fig. 9D)
reaching beyond end of scaphocerite; merus distinctly shorter
than carpus, 3.3 times as long as broad; carpus 1.2 times
as long as chela, 4.5 times as long as high; chela 2.5 times
as long as broad; fingers 1.5 times as long as palm. Third
pereiopod (Figs. 9E, F) reaching beyond end of scaphoceriteby length of dactylus, propodus 10 times as long as broad,
4.3 times as long as dactylus; dactylus 3.0 times as longas wide (spines included), with 5 or 6 accessory spines on
flexor margin. Fifth pereiopod (Figs. 9G, H) reaching beyond
end of scaphocerite, propodus 15 times as long as broad,3.0 times as long as dactylus, dactylus 3.5 times as long as
wide (spinules included), terminating in 1 claw, with 56
spinules on flexor margin.

Caridina valencia, new species(Fig. 9)Material examined. – Holotype: Male, cl 3.5 mm, USC, spring
Anislaq, outside, Valencia, coll. B. Sket, Feb.1995.
Paratypes: 5 males, cl 2.5–3.6 m, 92 juveniles, UOL, data same
as holotype.
Others: 1 juvenile, UOL, spring Badyang, cave, Valencia, coll.
B. Sket, Feb.1995; 1 male, cl 3.1 mm, 1 female, cl 2.8 mm, 1
female (damaged), 1 juvenile, UOL, Cave Castigio, Batuan, coll.
B. Sket, Feb.1995; 4 males, cl 2.2–2.4 mm, 15 females, cl 2.0–3.1
mm, 2 ovigerous females, cl 2.8–3.1 mm, ZRC 2007.0297, spring
below Cave Serpia, Ginguyuran, Dimiao, coll. B. Sket, Feb.1995;
1 male, cl 3.6 mm, 8 juveniles, ZRC 2007.0298, spring Anislaq,
cave, Valencia, coll. B. Sket, Feb.1995.

Endopod of male first pleopod (Fig. 9I) triangular, one-third
length of exopod, no appendix interna. Appendix masculina
of male second pleopod (Fig. 9J) half length of endopod,
with appendix interna half length of appendix masculina,
reaching near end of appendix masculina.

Description. – Rostrum (Fig. 9A) short, reaching near end of

Uropodal diaeresis (Fig. 9K) with 17 movable spinules.

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Fig. 9. Caridina valencia, new species: A, cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, first
pereiopod; D, second pereiopod; E, third pereiopod; F, dactylus of third pereiopod; G, fifth pereiopod; H, dactylus of fifth pereiopod; I,
endopod of male first pleopod; J, appendix masculina and appendix interna of male second pleopod; K, uropodal diaeresis. Scale bars: A= 1 mm; B, F, H = 0.1 mm; C–E, G = 0.5 mm; I–K = 0.2 mm. (males, cl 3.4 mm, paratype, ZRC, spring Anislaq, outside Cave Anislaq,
Valencia).

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Ovigerous females with egg sized 0.90 × 0.65 mm.

Incisor process of mandible ending in irregular teeth, molar
process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongated, with a number of distinct
teeth on inner margin, palp slender. Upper endites of maxilla
subdivided, palp short, scaphognathite tapering posteriorly
with some long, curved setae at posterior end. Palp offirst maxilliped rounded. Podobranch of second maxilliped
reduced to a lamina. Third maxilliped reaching to end of
scaphocerite, with ultimate segment as long as penultimate
segment.

Habitat. – Caridina valencia, new species, was collectedfrom several caves in the southern part of Bohol Island,
central Philippines.
Etymology. – The new species is named after its type localityValencia Town, Bohol Island, Philippines. The name is used
as a noun in apposition.
Remarks. – Caridina valencia, new species, morphologicallyresembles C. cebuensis Cai & Shokita, 2006a, from Cebu
Island, central Philippines. But it could be easily differentiatedfrom C. cebuensis by the much stouter carpus of the first
pereiopod (1.3 times as long as high vs. 1.6 times); theslender scaphocerite (3.6 times as long as broad vs. 3.1
times); and more spinules on the dactylus (56 vs. 39) (Fig.
9; cf. Cai & Shokita, 2006a: Figs. 1, 2).

Epipods on first 4 pereiopods. First pereiopod (Fig. 10D)
reaching end of antennular peduncle; merus 2.1 times aslong as broad, slightly shorter than carpus; carpus excavatedanteriorly, shorter than chela, 1.6 times as long as high;chela 2.1 times as long as broad; fingers slightly shorter thanpalm. Second pereiopod (Fig. 10E) reaching beyond end of
scaphocerite by length of chela; merus distinctly shorter thancarpus, 4.5 times as long as broad; carpus 1.3 times as longas chela, 5.3 times as long as high; chela 2.6 times as longas broad; fingers 1.3 times as long as palm. Third pereiopod(Figs. 10F, G) reaching beyond end of scaphocerite by lengthof dactylus and half of propodus, propodus 11 times as longas broad, 4.3 times as long as dactylus; dactylus 3.3 times aslong as wide (spines included), with 4 or 5 accessory spines
on flexor margin. Fifth pereiopod (Figs. 10H, I) reaching
beyond end of scaphocerite by length of dactylus, propodus 15
times as long as broad, 4.4 times as long as dactylus, dactylus
3.7 times as long as wide (spinules included), terminatingin 1 claw, with 44 spinules on flexor margin.

Caridina batuan, new species(Fig. 10)Material examined. – Holotype: Ovigerous female, cl 4.2 mm,
eggs 0.8 × 0.5 mm, USC, Cave Castigio, Batuan, coll. B. Sket,
Feb.1995.
Paratypes: One male, cl 4.1 mm, ZRC 2007.0299, same data as
holotype; 4 males, cl 2.2–3.7 mm, 5 females, cl 2.5–3.6 mm, 4juveniles, UOL, Cave Kalumpan, Behind-the-Clouds, Batuan, coll.
B. Sket, Feb.1995; 1 female, 1 juvenile, UOL, Spring Badyang,cave, Valencia, coll. B. Sket, Feb.1995; 2 females, cl 1.9–2.3 mm,
ZRC 2007.0300, Cave Bonugan, Batuan, coll. B. Sket, Feb.1995;
2 juveniles, UOL, Alijauan Bridge, stream, S. Duero, coll. B.
Sket, 5 Mar.1999.

Endopod of male first pleopod sub-triangular, a quarter lengthof exopod, no appendix interna. Appendix masculina ofmale second pleopod half length of endopod, with appendix
interna half length of appendix masculina, reaching near endof appendix masculina.

Description. – Rostrum (Fig 10A) short, reaching slightly
beyond end of basal segment of antennular pedunlcle;
rostral formula: 0–2+7–12/0–4. Antennal spine distinct,
fused with inferior orbital angle. Pterygostomian margin
broadly rounded.

Uropodal diaeresis (Fig. 10J) with 20 movable spinules.
Ovigerous females with eggs sized 0.80 × 0.50 mm.
Habitat. – Caridina batuan, new species, was collectedfrom several caves in southern part of the Bohol Island,
central Philippines.

Sixth abdominal somite 0.50 times of carapace, 1.9 times
as long as fifth somite, slightly shorter than telson. Telson(Fig. 10B, C) 2.9 times as long as wide, not terminating in a
projection, with 4 or 5 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of spines distinctly longer
than intermediate pairs. Preanal carina low, sub-triangular,
lacking spine.

Etymology. – The new species is named after its type localityBatuan, Bohol Island, the Philippines. The name is used asa noun in apposition.
Remarks. – Caridina batuan, new species, is morphologicallysimilar to C. minidentata Cai & Anker, 2004, from west
Samar, the Philippines. However, it can be distinguishedfrom C. minidentata by the more developed rostral teeth,
the much reduced eyes, the lower preanal carina and thelarger number of spinules on diaeresis (20 vs. 16) (Fig. 10;
cf. Cai & Anker, 2004: Figs. 4, 5).

Eyes reduced, anterior end reaching to 0.4 times length ofbasal segment of antennular peduncle. Antennular peduncle
0.63 times as long as carapace; basal segment of antennularpeduncle shorter than both second and third segment
lengths, anterolateral angle reaching 0.20 times length ofsecond segment, second segment distinctly longer than third
segment. Stylocerite reaching to 0.7 times length of basal
segment of antennular peduncle. Scaphocerite 3.4 times aslong as wide.

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Fig. 10. Caridina batuan, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D,
first pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fifth pereiopod; I, dactylus of fifth pereiopod;J, uropodal diaeresis. Scale bars: A, B = 1 mm; C = 0.2 mm, D–F, H = 0.5 mm; G, I, J = 0.1 mm (female, cl 3.0 mm, holotype, USC,
Cave Kalumpan, Batuan).

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Caridina anislaq, new species(Fig. 11)

to end of antennular peduncle, propodus 12 times as longas broad, 3.3 times as long as dactylus, dactylus 3.5 times
as long as wide (spinules included), terminating in 1 claw,
with 51 spinules on flexor margin.

Material examined. – Holotype: Male, cl 3.4 mm, USC.
Spring Anislaq, outside Cave Valencia, Valencia, coll. B. Sket,
Feb.1995.

Endopod of male first pleopod (Fig. 11J) triangular, one-third
length of exopod, no appendix interna. Appendix masculina(Fig. 11K) of male second pleopod half length of endopod,
with appendix interna half length of appendix masculina,
reaching near end of appendix masculina.

Paratypes: 10 males, cl 2.2–3.4 mm, 9 females, cl 2.2–5.4 mm,
ZRC 2007.0301, data same as holotype; 1 male, cl 2.4 mm, ZRC
2007.0302, Spring Anislaq, Cave Valencia, Valencia, coll. B.
Sket, Feb.1995.

Uropodal diaeresis (Fig. 11L) with 17 movable spinules.

Description. – Rostrum (Fig. 11A) short, reaching slightly
beyond end of basal segment of antennular pedunlcle; rostralformula: 3–6+6–11/1–6. Antennal spine fused with inferior
orbital angle. Pterygostomian margin broadly rounded.

Habitat. – Caridina anislaq, new species, was collectedfrom Spring Anislaq, near Cave Valencia, Valencia, Bohol
Island, central Philippines.

Sixth abdominal somite 0.48 times of carapace, 1.9 times
as long as fifth somite, slightly shorter than telson. Telson(Fig. 11B, C) 3.2 times as long as wide, not terminating ina projection, with 3 pairs of dorsal spinules and 1 pair of
dorsolateral spinules; lateral pair of distal spines distinctly
longer than intermediate pairs. Preanal carina low, subtriangular, lacking spine.

Etymology. – The new species is named after its type
locality-Spring Anislaq, Bohol Island, the Philippines, used
as a noun in apposition.
Remarks. – Caridina anislaq, new species, is morphologicallysimilar to C. batuan, new species, in the form of rostrum.
However, it could be easily separated from C. batuan by
the larger number of the post orbital rostral teeth (3–6 vs.
0–2), the stouter merus (1.2 times as long as broad vs. 2.1
times) and the carpus of the first pereiopod (as long as highvs. 1.6 times as long as high) and the shorter flagellum of
exopod of the first maxilliped (Figs. 10, 11).

Eyes well developed, anterior end reaching to 0.7 times
length of basal segment of antennular peduncle. Antennularpeduncle 0.70 times as long as carapace; basal segmentof antennular peduncle as long as both second and third
segment lengths, anterolateral angle reaching 0.20 times
length of second segment, second segment distinctly longer
than third segment. Stylocerite reaching to 0.7 times lengthof basal segment of antennular peduncle. Scaphocerite 3.6
times as long as wide.

Caridina camaro, new species(Fig. 12)Material examined. – Holotype: Female, cl 4.3 mm, USC, Cave
Camaro, Batuan, coll. B. Sket, Feb.1995.

Incisor process of mandible ending in irregular teeth, molar
process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongated, with a number of distinct
teeth on inner margin, palp slender. Upper endites of maxilla
subdivided, palp short, scaphognathite tapering posteriorly
with some long, curved setae at posterior end. Palp of first
maxilliped rounded, exopod with short flagellum. Podobranchof second maxilliped reduced to a lamina. Third maxillipedreaching beyond end of scaphocerite, with ultimate segment
slightly longer than penultimate segment.

Paratypes: Six more or less damaged specimens, ZRC 2007.0303,
data same as holotype.

Description. – Rostrum (Fig. 12A) short, reaching slightly
beyond end of basal segment of antennular pedunlcle; rostralformula: 2+10/5. Antennal spine fused with inferior orbital
angle. Pterygostomian margin broadly rounded.
Sixth abdominal somite 0.48 times of carapace, 1.8 times
as long as fifth somite, slightly shorter than telson. Telson(Figs. 12B, C) 2.7 times as long as wide, not terminating
in a projection, with 4 or 5 pairs of dorsal spinules and 1
pair of dorsolateral spinules; lateral pair of distal spines
distinctly longer than intermediate pairs. Preanal carina low,
sub-triangular, lacking spine.

Epipods on first 4 pereiopods. First pereiopod (Fig. 11D)
stout, reaching to middle of second segment of antennularpeduncle; merus 1.2 times as long as broad, slightly shorter
than carpus; carpus strongly excavated anteriorly, much
shorter than chela, as long as high; chela 2.1 times as long as
broad; fingers shorter than palm. Second pereiopod (Fig. 11E)
reaching to end of scaphocerite; merus distinctly shorter thancarpus, 3.3 times as long as broad; carpus slightly shorter or
as long as chela, 3.9 times as long as high; chela 2.4 times
as long as broad; fingers 1.5 times as long as palm. Third
pereiopod (Figs. 11F, G) reaching beyond end of scaphoceriteby length of 1/3 of propodus, propodus 12 times as longas broad, 4.9 times as long as dactylus; dactylus 2.5 times
as long as wide (spines included), with 4 accessory spines
on flexor margin. Fifth pereiopod (Figs. 11H, I) reachingEyes reduced, anterior end reaching to 0.4 times length ofbasal segment of antennular peduncle. Antennular peduncle
0.60 times as long as carapace; basal segment of antennularpeduncle distinctly as long as both second and third segment
lengths, anterolateral angle reaching 0.20 length of second
segment, second segment distinctly longer than third
segment. Stylocerite reaching to 0.7 times length of basal

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Fig. 11. Caridina anislaq, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D,
first pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fifth pereiopod; I, dactylus of fifth pereiopod;
J, endopod of male first pleopod; K, appendix masculina and appendix interna of male second pleopod; L, uropodal diaeresis. Scalebars: A, B = 1 mm; C = 0.2 mm, G, I = 0.1 mm; D–F, H, L = 0.5 mm (males, cl 3.4 mm, paratype, ZRC, spring Anislaq, outside Cave
Anislaq, Valencia).

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Fig. 12. Caridina camaro, new species: A, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D,
first pereiopod; E, second pereiopod; F, third pereiopod; G, dactylus of third pereiopod; H, fifth pereiopod; I, dactylus of fifth pereiopod;J, uropodal diaeresis. Scale bars: A, B = 1 mm; C = 0.2 mm; D–F, H = 0.5 mm; G, K = 0.1 mm; B, F, H = 0.1 mm; C–E, G = 0.5 mm;
I, J = 0.2 mm (females, cl 4.3 mm, holotype, USC, Cave Camaro, Batuan).

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segment of antennular peduncle. Scaphocerite 3.2 times aslong as wide.

Palaemonidae Rafinesque, 1815
Macrobrachium Bate, 1868

Incisor process of mandible ending in irregular teeth, molar
process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongate, with a number of distinct
teeth on inner margin, palp slender. Upper endites of maxilla
subdivided, palp short, scaphognathite tapering posteriorly
with some long, curved setae at posterior end. Palp of first
maxilliped rounded, exopod with short flagellum. Podobranchof second maxilliped reduced to a lamina. Third maxilliped
slender, with ultimate segment as long as penultimate
segment.

Macrobrachium rosenbergii rosenbergii (De Man, 1879)
Palaemon Rosenbergii De Man, 1879: 167 [type locality: Andai,
New Guinea (Irian Jaya), Indonesia].
Palamon carcinus rosenbergi – Ortmann, 1891: 701.
Palaemon (Eupalaemon) carcinus – De Man, 1902: 763 (not Cancer
carcinus Linnaeus, 1758).
Macrobrachium rosenbergii – Holthuis, 1950: 111 (part); Johnson,
1960: 260, Fig. 1; Chace & Bruce, 1993: 36, Fig. 15; Short,
2004: 44, Fig. 16 (part).
Macrobrachium rosenbergii rosenbergii – Johnson, 1973: 277;
Holthuis, 1995: 148; Cai & Ng, 2001: 674; Cai & Shokita,
2006a: 263.

Epipods on first 4 pereiopods. First pereiopod (Fig. 12D)
slender, merus 2.4 times as long as broad, shorter thancarpus; carpus excavated anteriorly, shorter than chela, 1.8 to
3.0 times as long as high; chela 2.7 times as long as broad;fingers 1.4 times as long as palm. Second pereiopod (Fig.
12E) elongated and slender, merus distinctly shorter thancarpus, 6.0 times as long as broad; carpus 1.5 times as longas chela, 7.7 times as long as high; chela 3.7 times as longas broad; fingers 1.9 times as long as palm. Third pereiopod(Figs. 12F, G) with propodus 16 times as long as broad, 4.4
times as long as dactylus; dactylus 5.6 times as long as wide
(spines included), with 6 accessory spines on flexor margin.
Fifth pereiopod (Figs. 12H, I) with propodus 15 times aslong as broad, 3.3 times as long as dactylus, dactylus 5 times
as long as wide (spinules included), terminating in 1 claw,
with 60 spinules on flexor margin.

Material examined. – One dried male, cl 103 mm, ZRC 2000.2407,
purchased from a fishermen who collected from the Loboc Riverin the 1980s Y. Cai, 19 Dec.2000; 1 molted cast, 2 juveniles, coll.
Y. Cai et al., Loboc River, Loboc, 19 Dec.2000.

Remarks. – Only one molted cast and two juveniles areavailable for the presnt study. Judging by the form of therostrum of the larger specimen, which is only slightly
beyond the end of scaphocerite and has a rostral formulaas 2+11/9. It apparently belongs to the eastern subspecies,
i.e. Macrobrachium rosenbergii rosenbergii.

Macrobrachium latidactylus (Thallwitz, 1891)

Uropodal diaeresis (Fig. 12J) with 17 movable spinules.

Palaemon latidactylus Thallwitz, 1891: 97 [type locality: Celebes(=Sulawesi), Indonesia]; Thallwitz, 1892:17, Fig. 3.
Palaemon (Macrobrachium) latidactylus – De Man: 1902: 805.
Palaemon lampropus De Man, 1892: 493, Pl. 29 Fig. 49 [typelocality: Celebes and Timor, Indonesia]; Kemp, 1918: 267.
Macrobrachium latidactylus – Holthuis, 1950: 239, Fig. 50; 1980:
97; Costa, 1979: 57; Naiyanetr, 1980: 17; Yeo et al., 1999:
236; Cai et al., 2004: 584; Short, 2004: 80, Fig. 31; Cai &Anker, 2004: 258; Wowor et al., 2004: 349, Fig 10E; Cai &Shokita, 2006a: 266.

Habitat. – Caridina camaro, new species, was collectedfrom the subterranean stream in Cave Camaro, Bohol Island,
central Philippines.
Etymology. – The new species is named after its type
locality—Cave Camaro, Bohol Island, the Philippines. The
name is used as a noun in apposition.
Remarks.- With regard to the slender pereiopods, Caridina
camaro, new species, is morphologically similar to C. liaoi,
new species. It can be easily differentiated from the latter
species by the reduced eyes, and the much shorter rostrum.
Compared with all the other cave species in Bohol, C. camaro
has more elongated carpus of the first pereiopods, hence,
could readily be distinguished from C. valencia, C. batuan
and C. anislaq (1.8–3.0 times as long as high vs. 1.0–1.6
times in the latter three species). It can be separated fromC. boholensis by the slender dactylus of third pereiopod (5.6
times as long as wide vs. 3.0 times)

Material examined. – 5 males, cl 7.3–15.1 mm, 5 females, cl
6.0–11.5 mm, ZRC 2007.0304, River outside Inabacan Cave,
Antequrra, coll. B. Sket, 23 Feb.1999; 1 male, cl 15 mm, ZRC
2007.0305, Alijanan Bridge, stream, S. Duero, coll. B. Sket, 5Mar.1999; 1 male, cl 10 mm, 1 female, cl 11 mm, ZRC 2007.0306,
River outside Inabacan Cave, Antequrra, coll. Y. Cai et al., 16
Dec.2000; 2 males, cl 12.8–13.5 mm, 1 female, cl 8.4 mm, ZRC
2007.0307, River outside Inabacan Cave, Antequrra, coll. Y. Cai
et al., 18 Dec.2000.

Remarks . – Macrobrachium latidactylus has been reported
from Sri Lanka (Costa, 1979), Indonesia, the Philippines,
Malaysia, southern Thailand (Holthuis, 1950; Johnson, 1963;
Chace & Bruce, 1993; Yeo et al., 1999), Hainan (Yu, 1936;
Liu et al., 1990), Taiwan (Hwang & Yu, 1982; Shy & Yu,
1998), the Ryukyu Islands (Shokita, 1979) and Australia(Short, 2004).

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Marobrachium australe (Guérin-Méneville, 1838)

Macrobrachium horstii (De Man, 1892)

Palaemon australis Guérin-Méneville, 1838: 37 [type locality:
Tahiti, French Polynesia].
Palaemon sundaicus Heller, 1862: 415, Pl. 2 Figs. 38, 39 [typelocality: Java, Indonesia].
Palaemon dispar Von Martens 1868: 41 [type locality: Pulau
Adonara, east of Flores, Indonesia].
Palaemon (Eupalaemon) dispar – De Man, 1902: 766.
Macrobrachium australis – Holthuis, 1950: 124, Figs. 27–30; Chace& Bruce, 1993: 23, Fig. 2; Cai & Ng, 2001: 683, Fig. 14a–d;
Cai & Anker, 2004: 258; Cai & Shokita, 2006a: 263.

Palaemon (Parapalaemon) Horstii De Man 1892: 460, Pl. 27: Fig.
39 [type locality: River at Polopo, central Celebes (Sulawesi),Indonesia].
Macrobrachium horstii - Holthuis, 1950: 203, Fig. 42; Chace &Bruce, 1993: 27; Cai & Ng, 2001: 679, Fig. 10; Wowor et al.,
2004: 349, Fig. 9L.
Material examined. – 1 male, cl 13.6 mm, 1 female, cl 16.8 mm,
1 ovigerous female, cl 12.0 mm, ZRC 2007.0311, River outside
Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male,
3 ovigerous females, cl 13.6 mm, ZRC 2007.0312, River outside
Inabacan Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000; 2 females,
cl 13.6, 6 ovigerous females, cl 11.0–13.8 mm, ZRC 2007.0313,
Loboc River, Loboc, coll. Y. Cai et al., 19 Dec.2000.

Material examined. – 3 males, cl 17.0–24 mm, 6 ovigerous females,
15.7–19.7 mm, ZRC 2007.0308, Loboc River, Loboc, coll Y. Cai
et al., 19 Dec.2000.

Remarks. – Macrobrachium horstii has previously beenreported from Leyte, the Philippines (Cai & Ng, 2001).
It is currently known from the Philippines and eastern
Indonesia.

Remarks. – The species has a wide distribution in the
Indo-West Pacific, from Madagascar to Polynesia (Holthuis,
1950; Chace & Bruce, 1993).

Macrobrachium lar (Fabricius, 1798)

Macrobrachium placidulum (De Man, 1892)

Palaemon Lar Weber, 1795: 94 [nomen nudum]
Palaemon Lar Fabricius, 1798: 402 [type locality: in India orientali
Dom. Daldorf]; Cowles, 1914: 380, Pl. 2 Fig. 7.
Palaemon (Eupalaemon) lar – De Man, 1902: 774.
Macrobrachium lar – Holthuis, 1950: 176, Fig. 37; Chace & Bruce,
1993: 30, Fig. 9; Yeo et al., 1999: 236; Cai & Ng, 2001: 683,
Fig. 14e; Wowor & Choy, 2001: 285; Cai & Ng, 2002: 75;
Cai & Anker, 2004: 256; Cai et al., 2004: 585; Wowor et al.,
2004: 353, Fig. 14; Short, 2004: 75, Fig. 29.

Palaemon (Macrobrachium) placidulus De Man, 1892: 489, Pl.
28 Fig. 48 [type localities: Celebes (Sulawesi), Pulau Selajar,
Flores, and Timor].
Macrobrachium placidulum – Holthuis, 1950: 253, Fig. 51c; Chace& Bruce, 1993: 35, Fig. 14; Cai & Shokita, 2006a: 264.
Macrobrachium placidum – Shokita, 1979: 275.
Macrobrachium horstii – Hwang & Yu, 1982: 164, Fig. 5 (notPalaemon (Parapalaemon) Horstii De Man 1892).
Macrobrachium cf horstii – Shy & Yu, 1998: 30.

Material examined. – One male, cl 22.6 mm, ZRC 2007.0309,
Loboc River, Loboc, coll. Y. Cai et al., 19 Dec.2000.

Material examined. – One male, cl 14.2 m, ZRC 2007.0314,
Alijanan Bridge, stream, S. Duero, coll. B. Sket, 5 Mar.1999; 3
males, cl 15.3–17.4 mm, ZRC 2007.0315, river outside Inabacan
Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male, cl 8.4
mm, 1 female, cl 10.3 mm, ZRC 2007.0316, river outside Inabacan
Cave, Antequrra, coll. Y. Cai et al., 18 Dec.2000.

Remarks. – Macrobrachium lar is a well known species,
widely distributed throughout the Indo-West Pacific.
However, it seems rather rare in Bohol.

Remarks. – Macrobrachium placidulum is a common
freshwater species in the Philippines. It is distributed from
Ryukyu Islands, Taiwan, the Philippines, Palau, and eastern
Indonesia, to New Guinea, as well as Fiji (Chace & Bruce,
1993; Choy, 1984).

Macrobrachium scabriculum (Heller, 1862)
Palaemon scabriculus Heller, 1862: 527 [type locality: Sri Lanka].Henderson & Matthai, 1910: 296, Pl.17 Figs. 7a–c, Pl. 18
Figs. 7a–p.
Macrobrachium scabriculum – Holthuis, 1950: 224; Chace & Bruce,
1993: 37; Johnson, 1973: 15; Yeo et al., 1999: 231, Figs. 18; 19;
Wowor & Choy, 2001: 286; Cai & Ng, 2002: Fig. 16; Wowor
et al., 2004: 350, Fig. 12I; Cai & Shokita, 2006a: 266.

Macrobcrachium equidens (Dana, 1852)
Palaemon equidens Dana, 1852: 26 [type locality: Singapore].
Palaemon sundaicus – Cowles, 1914: 355, Pl. 2 Fig. 3 (not Palaemon
sundaicus Heller, 1862).
Macrobrachium equidens – Holthuis, 1950:162, Fig. 36; 1980: 90;
Liu et al., 1990: 110, Fig. 8; Chace & Bruce, 1993: 25, Fig.
4; Yeo et al., 1999: 226; Wowor & Choy, 2001: 282; Cai &Anker, 2004: 389; Cai et al., 2004: 589; Wowor et al., 2004:
349, Fig. 10H; Short, 2004: 26, Figs. 8, 9; Cai & Shokita,
2006a: 265.

Material examined. – One male, cl 19.5 mm, 1 ovigerous female,
24 mm, ZRC 2007.0310, Loboc River, Loboc, coll. Y. Cai et al.,19 Dec.2000.

Remarks. – Macrobrachium scabriculum was reported
recently from Mindanao (Cai & Shokita, 2006a). This is thesecond record for the species to occur in the Philippines.

Material examined. – 1 male, cl 7.5 mm, ZRC, River outside
Inabacan Cave, Antequrra, coll. Y. Cai et al., 16 Dec.2000; 1 male,cl 10.5 mm, 1 female, cl 10.0 mm, ZRC, Loboc River, Loboc, coll.
Y. Cai et al., 19 Dec.2000.

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Bouvier, E. L., 1925. Recherches sur la morphologie, les variations,la distribution géographique des crevettes des la famille desAtyidés. Encyclopédie Entomologique, series A, 4: 1–370,
Figs. 1–761.

Remarks. – Marcobrachium equidens is a brackish water
species, with gravid females and smaller specimens being
commonly found in mangrove creeks. It is known from avery wide area in the Indo-West Pacific, from Madagascar to
the Solomon Islands and Fiji (Choy, 1984; Chace & Bruce,
1993; Short, 2004). However, a recent molecular analysis
results (Liu et al., 2007) show that the incongruence of
non-monophyly of M. equidens, in 16S, COI and 16S+COI
analyses, implied the existence of a cryptic species in thePhilippines and Taiwan.

Cai, Y. & A. Anker, 2004. On a collection of freshwater shrimps
from the Philippines, with descriptions of five new species.
Tropical Zoolology, 17: 233–266.
Cai, Y., P. Naiyanetr & P. K. L. Ng, 2004. The freshwater
prawns of the genus Macrobrachium Bate, 1868, of Thailand(Crustacea: Decapoda: Palaemonidae). Journal of Natural
History, 38: 581–649.
Cai, Y., & P. K. L., Ng, 2001. The freshwater decapod crustaceansof Halmahera, Indonesia. Journal of Crustacean Biology, 21:
665–695, Figs. 1–19.

Palaemon Weber, 1795

Cai, Y. & P. K. L. Ng, 2002. The freshwater palaemonid prawns(Crustacea: Decapoda: Caridea) of Myanmar. Hydrobiologia,
487: 59–83.

Palaemon concinnus Dana, 1852
Palaemon concinnus Dana, 1852: 587 [type locality: Fiji Islands];
Holthuis, 1950: 61, Fig. 12; Chace & Bruce, 1993: 40; Cai &Ng, 2001: 686, Fig. 14f; Cai & Shokita, 2006a: 267.

Cai, Y. & P. K. L. Ng, 2007. A revision of the Caridina gracilirostris
De Man, 1892, species group, with descriptions of two new
taxa (Decapoda; Caridea; Atyidae). Journal of Natural History,
41(25–28): 1585–1602.

Material examined. – 2 males, cl 7.0–7.6 mm, 1 female, cl 9.2mm, ZRC 2007.0317, tributary of Loboc River, Loboc, coll. Y.
Cai et al., 19 Dec.2000.

Cai, Y. & S. Shokita, 2006a. Report on a collection of freshwater
shrimps (Crustacea: Decapoda: Caridea) from the Philippines,
with descriptions of four new species. Raffles Bulletin of
Zoology, 54: 245–270.

Remarks. – Palaemon concinnus has a wide distribution
in the Indo-West Pacific, from South Africa, Indonesia,
Philippines to Marshall Islands and Tuamotu Archipelago
in French Polynesia (Chace & Bruce, 1993). The species
is commonly found in brackish to fresh water in the lower
reaches of rivers.

Cai, Y., & S, Shokita, 2006b. Atyid shrimps of the Ryukyu Islands,
Southern Japan (Crustacea: Decapoda: Atyidae). Journal of
Natural History, 40(38–40): 2123–2172.
Cai, Y., P. K. L. Ng, S. Shokita & K. Satake, 2006. Revision of
the Japanese atyid shrimps (Decapoda: Caridea) first describedby William Stimpson, 1860. Journal of Crustacean Biology,
26(3): 392–419.

ACKNOWLEDGEMENTS

Chace, F. A. Jr., 1983. The Atya-like shrimps of the Indo-pacificRegion (Decapoda: Atyidae). Smithsonian Contributions to
Zoology, 384: 1–54, 24 figures.

The first author would like to thank Boris Sket (UOL) for
sending his specimens and the accompanying habitat notes.
The ZRC-USC Expedition was led by Lawrence Liao (USC),
and we are grateful to the support cum help of Benito Tan
and N. K. Ng (ZRC), and Adonis Porpetcho and Roxie Diaz
(USC). The Caving Team from the Holy Name University
at Bohol was instrumental in helping with the speleologicalsurveys, and the support of their president, Florentes Camacho
is gratefully acknowledged. Boris Sket read the manuscript
and to N. K. Ng kindly prepared the map.

Chace, F. A., Jr., 1997. The Caridean shrimps (Crustacea:
Decapoda) of the Albatross Philippine expedition 1907–1910.
Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae,
Bathypalaemonellidae, Processidae, and Hippolytidae.
Smithsonian Contributions to Zoology, 58: 1–106, Figs.1–29.
Chace, F. A. Jr. & A. J. Bruce, 1993. The Caridean shrimps(Crustacea: Decapoda) of the Albatross Philippine expedition
1907–1910. Part 6: Superfamily Palaemonoidea. Smithsonian
Contributions to Zoology, 543: i–vii + 1–152, Figs. 1–23.
Choy, S., 1984. On the freshwater palaemonid prawns from the Fiji
Islands (Decapoda, Caridea). Crustaceana, 47: 269–277.

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