JOURNAL OF CRUSTACEAN BIOLOGY, 26(3): 392–419, 2006
ON THE SPECIES OF JAPANESE ATYID SHRIMPS (DECAPODA: CARIDEA)
DESCRIBED BY WILLIAM STIMPSON (1860)
Yixiong Cai, Peter K. L. Ng, Shigemitsu Shokita, and Kiyoshi Satake
(YC) Tropical Marine Science Institute, National University of Singapore, 14 Kent Ridge Road,
Singapore 119223, Republic of Singapore (caiyixiong@yahoo.com);
(PKLN) Department of Biological Sciences, National University of Singapore, Lower Kent Ridge Road,
Singapore 119260, Republic of Singapore (dbsngkl@nus.edu.sg);
(SS) Department of Chemistry, Biology and Marine Science, Faculty of Science, University of the Ryukyus,
Nishihara, Okinawa 903-0213, Japan (shokita@sci.u-ryukyu.ac.jp);
(KS) Division of Environmental Biology, National Institute for Environmental Studies,
Onogawa, Tsukuba, Ibaraki 305-8506, Japan (satanii@nies.go.jp)
ABSTRACT
The taxonomic status of the six Japanese species of Caridina described by W. Stimpson (1860) is clarified on the basis of fresh specimens
from the type localities. Caridina grandirostris, which has long been synonymized under C. longirostris H. Milne Edwards, 1837, is
shown to be a distinct species; C. leucosticta is redescribed; C. multidentata is the senior synonym of C. japonica De Man, 1892; C.
acuminata and C. brevirostris are regarded as junior synonyms of Atyoida pilipes (Newport, 1847); and C. exilirostris is synonymized
with C. typus H. Milne Edwards, 1837. Neotypes for the six species are designated to stabilize their taxonomy, all of which are redescribed
and figured. The various nomenclatural problems associated with these species are discussed.
INTRODUCTION
On the basis of the United States North Pacific Exploring
Expedition (see Habersham, 1857, for details of the cruise),
Stimpson (1860) published preliminary descriptions of
seven new species of atyid shrimps of the genus Caridina:
C. grandirostris, C. leucosticta, C. multidentata, C. serrata,
C. acuminata, C. brevirostris and C. exilirostris (see also
Stimpson, 1907). Unfortunately, almost the entire crustacean
collection together with most of the associated manuscripts
in preparation and drawings were lost when the Chicago
Academy of Sciences was destroyed in the great fire of
1871 (see Deiss and Manning, 1981). Although some of
Stimpson’s syntypes have subsequently been found in the
British Museum (Natural History), London, no species of
Caridina are known to be extant (Evan, 1967). The
descriptions of the new species of Caridina in Stimpson
(1860) were very brief and no figures provided. This has
caused many taxonomic and nomenclature problems over the
years. One of these ‘‘problematic’’ taxa, Caridina serrata,
originally described from Hong Kong, was redescribed in
detail by Cai and Ng (1999) and a neotype was designated.
In the present study, the taxonomic status of the
remaining six Japanese species of Caridina described by
Stimpson (1860) are clarified on the basis of a large number
of fresh specimens from the type localities. Redescriptions
are provided, the species are illustrated in detail, and the
various taxonomic and nomenclature problems involved are
discussed. Specimens involved in this study are deposited
in the National Science Museum, Tokyo, Japan (NSMT);
Department of Chemistry, Biology and Marine Science,
Faculty of Science, University of the Ryukyus, Okinawa,
Japan (UR); National Institute for Environmental Studies,
Ibaraki, Japan (NIES); Zoological Reference Collection,
Raffles Museum of Biodiversity Research, National University of Singapore, Singapore (ZRC); National Museum
of Natural History, Leiden, the Netherlands (RMNH);
Zoological Museum, Amsterdam, the Netherlands (ZMA);
Basel Natural History Museum, Basel, Switzerland
(NHMB); Senckenberg Museum, Frankfurt am Main; Germany (SMF); and Muséum National d’Historie Naturelle,
Paris, France (MNHN). The abbreviation cl is used for
carapace length, measured (in millimeters) from the postorbital margin to the posterior dorsal margin of the carapace.
Notation for rostral formula follows that of Chace and
Bruce (1993).
SYSTEMATICS
Atyoida pilipes (Newport, 1847)
Figs. 1-3
Atya pilipes Newport, 1847:160 [type locality: Apia, Upoln, Navigator or
Samoan group (see Chace, 1983)].
Atya pilipes: A. Milne-Edwards, 1864:150; De Man, 1892a: 363; Bouvier,
1925: 304; Shokita, 1979: 199.
Atyoida tahitensis Stimpson, 1860: 28 [type locality: Tahiti, French
Polynesia].
Atyoida pilipes: Smith and Williams, 1982: 345 (part); Chace, 1983: 10,
figures 3-8; Chace, 1997: 4; Hayashi, 1989a: 127, fig. 161; Shokita,
1997: 507.
Caridina acuminata Stimpson, 1860: 98 [type locality: Ogasawara (Bonin)
Islands, Japan]; Ortmann, 1895: 401; Balss, 1914: 24, fig. 18; Kamita,
1976:23.
Caridina typus acuminata: Bouvier, 1925: 252 (part).
Not Caridina acuminata: Shokita, 1979: 205; Hayashi, 1989c: 311, fig.
169b, c, 170f.
Not Caridina typus acuminata: Bouvier, 1925: 126, figures 271.
Not Caridina typa acuminata: Roux, 1934: 222.
Caridina brevirostris Stimpson, 1860: 98 [type locality: Okinawa (Loo
Choo) Island, Ryukyu Islands, Japan]; Bouvier, 1905: 78 (part).
Not Caridina brevirostris: Bouvier, 1904: 136; Rathbun, 1906: 919, fig. 67;
Edmondson, 1935: 16; Fujino, 1972: 7, fig. 5.
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CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
393
Fig. 1. Atyoida pilipes. A, cephalothorax and cephalic appendages, lateral view; B, preanal carina; C, telson, D, mandible; E, maxillula; F, maxilla;
G, first maxilliped; H, second maxilliped; I, third maxilliped; J, scaphocerite. Scales: A, I ¼ 1.0 mm; B-H, J ¼ 0.5 mm. (neotype of Caridina acuminata,
male, cl 5.7 mm, NSMT, Chichi-jima Island)
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Fig. 2. Atyoida pilipes. A, first pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fourth pereiopod; F, dactylus of fourth
pereiopod; G, fifth pereiopod; H, dactylus of fifth pereiopod. Scales: A, B ¼ 1.0 mm; C, E, G ¼ 0.5 mm; D, F, H ¼ 0.5 mm; G, I ¼ 0.2 mm. (neotype of
Caridina acuminata, cl 5.7 mm, NSMT, Chichi-jima Island)
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
395
Fig. 3. Atyoida pilipes. A, male first pleopod; B, endopod of male first pleopod; C, male second pereiopod; D, appendix masculina ans appendix interna
of male second pleopod; E, distal portion of telson; F, uropodal diaeresis. Scales: A, C ¼ 0.5 mm; B, D, E, F ¼ 0.2 mm. (neotype of Caridina acuminata,
cl 5.7 mm, NSMT, Chichi-jima Island)
Material Examined.—Neotype of Caridina acuminata: 1
male, cl 6.5 mm, NSMT, a stream of Naka-kaigan Coast,
lotic system, EC 380, 60 to 70 meters above sea level,
Chichi-jima Island, Ogasawara Islands, coll. K. Satake, R.
Ueno and M. Inaba, 28 Feb 2000. Neotype of Caridina
brevirostris: 1 male, cl 6.8 mm, NSMT, River Aritsu,
Kume-jima Island, coll. A. Kawakami, 8 Dec 1995. Others:
6 males, cl 4.1-5.8 mm, NIES-H-3, Nagahama-bashi, lotic
system, EC 270-610, 130 to 140 meters above sea level,
Haha-jima Island, Ogasawara Islands, coll. K. Satake, 13
Feb 1998; 5 males, cl 5.3-6.6 mm, 14 females, cl 4.5-8.6
mm; 2 ovigerous females, cl 8.3-8.8 mm, NIES-H-6, upper
reach of Koromodate Stream, freshwater, lotic system, EC
590-700, 70 to 110 meters above sea level, Haha-jima Is-
land, Ogasawara Islands, coll. K. Satake and R. Ueno, 24
Feb 2000; 10 females, cl 7.5-8.9 mm (3 females, ZRC
2004.0518), NIES-CH-17, a stream of Naka-kaigan Coast;
freshwater; lotic system; EC 380; 60 to 70 meters above sea
level, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake, R. Ueno and M. Inaba, 28 Feb 2000; 1 male, cl
5.4 mm, 4 females, cl 6.8-7.4 mm; NIES-CH-23, a stream of
Tenno-ura, lotic system, EC 610, 90 meters above sea level,
Chichi-jima Island, Ogasawara Islands, coll. K. Satake and
H. Suzuki, 4 May 2000; 2 males, cl 6.2-7.1 mm, UR, Omija
River, Iriomote, coll. T. Naruse, 1 Aug 1999.
Description.—Rostrum short, with dorsal margin convex,
apex slightly directed ventrally, reaching near to or slightly
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beyond end of basal segment of antennular peduncle,
median dorsal carina unarmed, ventral margin unarmed,
rarely with 1 or 2 teeth; antennal spine fused with inferior
orbital angle, acute; pterygostomian angle acute. Sixth
abdominal somite 0.4 times as long as carapace, 1.3 times as
long as fifth somite, shorter than telson. Telson 3.0 times as
long as wide, with 5-7 pairs (or 10-12 if not in pairs) dorsal
spinules, situated on distal half, 1 pair of dorsolateral spines
near distal end, posterior margin with 4 inconspicuous
median fixed teeth and 5 pairs of spine-like plumose setae
on distal margin, lateral pair distinctly shorter than intermediate pairs; preanal carina triangular, without spine.
Eyes well developed. Antennular peduncle stout, half
length of carapace; basal segment shorter than half length
of antennular peduncle, second segment longer than third
segment; stylocerite reaching slightly beyond end of basal
segment. Scaphocerite 2.5 times as long as wide, inner
margin straight, with large distolateral tooth reaching near
end of antennular peduncle.
Incisor process of mandible ending in irregular teeth,
molar process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongate, widened distally, with
many distinct teeth on inner margin, palp slender. Upper
endites of maxilla subdivided, palp short, scaphognathite
tapering posteriorly with more than 20 long setae at posterior end. Palp of first maxilliped ending in a finger-like
process. Second maxilliped with ultimate segment not fused
with penutimate segment; well developed podobranch present. Third maxilliped reaching near end of antennular peduncle, with a short exopod, reaching only slightly beyond
end of antepenutimate segment, ultimate segment as long as
penultimate segment.
Epipods on first 4 pereiopods. First pereiopod short,
reaching to end of basal segment of antennular peduncle,
merus 1.5 times as long as broad; carpus excavated strongly
anteriorly, length shorter than width; chela variable, with
almost no discernable palm to palm longer than finger. Second
pereiopod reaching end of proximal segment of antennular
peduncle, merus more slender than that of first pereiopod,
more than twice as long as width, carpus excavated strongly
anteriorly, as long as width or shorter than width, chela
variable, similar to previous leg. Last 3 pereiopods subequal
in length. Third pereiopod with row of setae on outer surface
of each joint, reaching to end of antennular peduncle,
propodus distinctly shorter than merus, 5.6 times as long as
broad, 3.0 times as long as dactylus; dactylus ending in
a strong claw; 2.5 times as long as wide (spines included),
flexor margin with 4 accessory spines. Fourth pereiopod
reaching slightly beyond end of proximal segment of antennular peduncle, similar to third pereiopod in form. Fifth
pereiopod, with setae only present on merus, reaching slightly
beyond post orbital margin, propodus longer than merus, 8.5
times as long as broad, 3.2 times as long as dactylus; dactylus
4.2 times as long as wide, with 6-10 spinules on flex or margin.
Endopod of male first pleopod extending to half length of
exopod, subtriangular, 2.6 times as long as broad, with
a distinct appendix interna near distal end of endopod.
Appendix masculina of male second pleopod slender,
reaching to 2/3 length of endopod, inner and distal surface
densely lined with long spines; appendix interna at basal 2/5
of appendix masculina, extending to distal 1/3 of appendix
masculina. Diaeresis of uropodal exopod with 19-23
spinules.
Remarks.—Stimpson (1860) gave a very short description
of C. acuminata: ‘‘Thorax sat compressus. Rostrum breve,
oculos parce superans, trigonum, ad basin horizontaliter
latum, ad extremitate paullo deflexum; marginibus totis
levibus; crista dorsali non dilatata, dorso continua. Antennularum flagella longitudine aequalia. Manuum penicilli
parvi, breves. Pedes postici spinulis asperi; tertii et quinti
paris quam quarti paris longiores. Color olivaceus, punctatus.’’ [Carapace compressed. Rostrum short, situated above
eyes, triangular, horizontal at base laterally; tip slightly
downturned, margin normally smooth, crest of dorsal
margin of rostrum not dilated. Antennular flagella as long
as body length. Chela short, with setae. Walking legs armed
with rough spinules; third and fifth legs as long as fourth
leg. Olivaceous in colour, spotted.]
During a recent faunistic survey of the Ogasawara Islands
(¼ Bonin Islands), the type locality of C. acuminata, six
species of atyid shrimps were found, including three species
of Caridina, namely C. multidentata Stimpson, 1860 (¼ C.
japonica De Man, 1892b, see later), C. typus H. Milne
Edwards, 1837, C. celebensis De Man, 1892, one species of
Atyoida and one species of Paratya (see Satake and Cai,
2005). Utilising the specimens on hand and reading the brief
description of Stimpson (1860) carefully, it leaves us in little
doubt that C. acuminata closely matches what is today
called Atyoida pilipes. It certainly does not appear to be
a species of Caridina.
Balss (1914) recorded C. acuminata from Ito, Japan.
Kubo (1938), however, commented that Balss’ specimens
‘‘. . . show discrepancy to the material at my disposal in
having 7 spinules (instead of 60 setae) on the posterior
border of fifth leg.’’ It seems to us that Balss’ specimens
are also Atyoida pilipes while Kubo’s material probably
belongs to C. typus instead. The exact identities of their
shrimps, however, can only be ascertained after this material
is reexamined. Bouvier (1925: 126, figure 271) provided
detailed illustrations of Caridina typus var. acuminata, on
the basis of specimens from Poulo Condor (So’n Island),
southern Vietnam. His specimens are clearly referable to C.
typus H. Milne Edwards, 1837 (see remarks under C. typus)
instead. The C. typa acuminata from Admiralty Islands
reported by Roux (1934) should also be assigned to C.
typus. Kamita (1976) reported C. acuminata from Hawai’i.
According to his description and figures, notably in the form
of rostrum, stout carpus of second pereiopod (1.5 times as
long as high), and number of spinules on the dactylus of the
fifth pereiopod (15), his species is almost certainly A.
bisulcata Randall, 1840, instead. Although A. bisulcata is
very close to A. pilipes and cannot be separated by
characters given by Kamita (1976), A. pilipes is not known
from the Hawaiian Islands as yet (Eldredge and Miller,
1997). Shokita (1979) reported C. acuminata from Arakawa
River, Okinawa. These specimens are no longer extant, but
fresh material collected from the same locality shows that
C. typus is the only species there, and it seems a fair
assumption that Shokita’s record of C. acuminata is almost
certainly C. typus instead.
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
A second species described by Stimpson (1860) is also
involved in this confusion. Stimpson (1860) described C.
brevirostris in a short passage: ‘‘Corpus gracile. Rostrum
brevissimum, oculis brevius, trigonum; margine superiore
obtuso, laevi. Margo carapacis ad basin antennarum inermis.
Manus primi paris digiti breves, quam palma multo breviores.
Pedum posticorum dactyli robusti, vix curvati; et quartam
partem articuli penultimi;(?) longitudine aequantes. Long 0.5
poll. C. acuminata affinis, rostrobreviore. Hab.-Ad insulam
‘‘Loo Choo’’; in aquis dulcibus.’’ [Body slender. Rostrum
very short, unarmed, not reaching beyond eye stalk,
triangular. Margin of carapace reaching base of antenna.
Chela of first pair of legs with short fingers, much shorter than
palm. Dactylus of posterior legs stout, very curved and 1/4 as
long as propodus. Short rostrum similar to C. acuminata.]
The type locality, Loo Choo Island, is now called Okinawa
(T. Naruse, personal communication). This description, albeit
brief, actually agrees well with young specimens of Atyoida
pilipes. Atyoida pilipes has been reported from Yona River of
Okinawa Island by Shokita (1979). Unfortunately, Shokita’s
(1979) specimens are no longer extant, and as recent collections from the Yona River did not reveal any specimens
of A. pilipes, his identification cannot be confirmed.
Bouvier (1904: 136; 1925: 227) reported C. brevirostris
from the Seychelles. His material (56 specimens, MNHN
Na 690-693) was re-examined by the first author. These
specimens have a variable rostrum with ovigerous females
having large eggs, and they most likely belong to an
undescribed species of Caridina. They certainly do not match
what Stimpson (1860) described as C. brevirostris. The
‘‘Caridina brevirostris’’ reported by Rathbun (1906: 919,
figure 67) and Edmondson (1935: 16) from the Hawaiian
Islands has been referred to Halocaridina rubra Holthuis,
1963 (cf. Holthuis, 1973). Kubo (1941) reported and
described in detail specimens he identified as C. brevirostris
from Ishigaki Island, and transferred the species to
Neocaridina Kubo, 1938. This record has been cited by
several subsequent authors (Shokita, 1975; 1979; Miyake,
1977; Hayashi, 1990; Cai, 1996). As discussed above, the
true C. brevirostris Stimpson, 1860, is now regarded as a
junior synonym of Atyoida pilipes. Kubo’s specimens are
no longer extant in the Tokyo University of Fisheries, where
the late Prof. Itsuo Kubo worked (T. Naruse, personal
communication), and subsequent surveys on the freshwater
shrimp fauna of Ishigaki did not obtain any specimens that
can be referred to Kubo’s ‘‘N. brevirostris.’’ ‘‘Neocaridina
brevirostris’’ has also been reported from Iriomote Island
(Shokita, 1975; 1979). Reexamination of specimens from
Iriomote Island has shown that they belong to a new species
of Neocaridina instead (Naruse et al., 2006). For the
moment, Kubo’s (1941) specimens are referred to an
undescribed species of Neocaridina Kubo, 1938.
To prevent further taxonomic confusion with the identities of Stimpson’s (1860) species, we here designate an
adult male specimen (cl 6.5 mm, NSMT) of Atyoida pilipes
from Chichi-jima Island, Ogasawara Islands, as the neotype
of Caridina acuminata Stimpson, 1860; and one male (cl
6.8 mm, NSMT), from Kume-jima Island, a small island
near Okinawa Island, as the neotype of C. brevirostris
Stimpson, 1860.
397
Distribution.—Atyoida pilipes has been reported from the
Philippines and eastern Lesser Sunda Islands at about
1208E, and eastward through the Pacific high islands
(Chace, 1983). In Japan, it has been reported from Okinawa,
Ishigaki (Shokita, 1979), Iriomote (Shokita, 1997) and
Kume-jima Island of the Ryukyus.
Habitat.—Atyoida pilipes lives among the roots of trees or
vegetation of fast flowing waters. It also occurs in small
waterfalls with vertical trenches that serve as hiding places.
Caridina grandirostris Stimpson, 1860
Figs. 4-6
Caridina grandirostris Stimpson, 1860: 97 [type locality: Okinawa (Loo
Choo) Island, Ryukyus, Japan].
Caridina grandirostris: Kubo, 1938: 85, figs. 15a-m.
Caridina nilotica: Fujino, 1972: 7; Miyake, 1977: 39; Shokita, 1975: 119;
Caridina longirostris: Shokita, 1979: 202; Hayashi, 1989c: 312, fig. 169e;
Suzuki et al., 1993: 56, Suzuki and Sato, 1994: 62.
Material Examined.—Neotype: 1 male, cl 4.9 mm, NSMT,
upper stream of Tima River, about 4 km from river mouth,
Okinawa Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng, T.
Naruse and S. Islam, 11 Jun 2000. Others: 3 males, cl 3.23.8 mm, 1 female, cl 4.3 mm, 6 ovigerous females, cl 4.25.3 mm, UR, 26833.489N 128802.489E, freshwater stream
draining to a small patch of mangrove, Oura River, Okinawa
Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng, T. Naruse
and S. Islam, 11 Jun 2000; 13 ovigerous females, cl 4.6-5.6
mm, 4 females, cl 3.6-4.6 mm, 10 males, cl 3.1-3.8 mm,
ZRC 2004.0519, 26833.429N 128804.609E, upstream of
Tima River, about 1-2 km from river mouth, Okinawa
Island, Ryukyu Islands, pH 7.4, coll. Y. Cai, N. K. Ng, T.
Naruse and S. Islam, 11 Jun 2000; 46 males, cl 3.6-4.6 mm,
12 females, cl 3.6-4.9 mm, 1 ovigerous female, cl 4.7 mm,
ZRC 2004.0520, 26833.639N 128805.529E, upstream of
Tima River, about 3 Km from the river mouth, Okinawa
Island, Ryukyu Islands, pH 7.4, coll. N. K. Ng and S. Islam,
11 Jun 2000; 6 males, cl 3.8-5.1 mm, 12 females, cl 4.7-6.5
mm, 4 ovigerous females, cl 5.8-6.5 mm, ZRC 2004.0521,
upper stream of Tima River, about 4 km from river mouth,
Okinawa Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng, T.
Naruse and S. Islam, 11 Jun 2000; 20 males, cl 3.5-4.4 mm,
1 female, cl 5.2 mm, 25 ovigerous females, cl 5.7-6.1 mm,
ZRC 2004.0522, 26836.619N 128807.049E, Arume River,
Okinawa Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng, T.
Naruse and S. Islam, 11 Jun 2000; 8 males, cl 2.9-4.0 mm,
10 females, cl 5.0-5.6 mm, 15 ovigerous females, cl 5.0-5.6
mm, ZRC 2004.0523, 24830.119N 124815.269E, Gaburumata River, Ishigaki Island, Ryukyu Islands, pH 7.3, coll. Y.
Cai and T. Naruse, 13 Jun 2000; 2 males, cl 3.6-4.1 mm, 1
female, cl 6.8 mm, 1 ovigerous female, cl 6.1 mm, ZRC
2004.0524, 24824.429N 124809.809E, fast flowing stream,
with pH 7.6 at one of the tributaries of Nagura River, below
reservoir, Ishigaki Island, Ryukyu Islands, coll. Y. Cai and
T. Naruse, 13 Jun 2000; 2 males, cl 3.9-4.0 mm, 1 female, cl
5.6 mm, 1 ovigerous female, cl 5.3 mm, ZRC 2004.5025,
24825.009N 124809.869E, fast flowing tributary of Nagura
River, below a reservoir, Ishigaki Island, Ryukyu Islands,
pH 7.2, coll. Y. Cai and T. Naruse, 13 Jun 2000; 1 female, cl
6.1 mm, ZRC 2004.5026, 24823.659N 123851.849E, fast
flowing water, about 200 metres from sea, Omija River,
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 26, NO. 3, 2006
Fig. 4. Caridina grandirostris. A, cephalothorax and cephalic appendages, lateral view; B, preanal carina; C, telson, D, distal portion of telson,
E, scaphocerite; F, mandible; G, maxillula; H, maxilla; I, first maxilliped; J, second maxilliped; K, third maxilliped; L, uropodal diaeresis. Scales: A, C ¼
1.0 mm; B, E-K ¼ 0.5 mm, D, L ¼ 0.2 mm. (neotype male, cl 4.9 mm, NSMT, Tima River, Okinawa, Ryukyu Islands)
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
Iriomote Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng and
T. Naruse, 15 Jun 2000; 1 male, cl 4.0 mm, ZRC 2004.0527,
fast flowing water, upper part of Omija River, Iriomote
Island, Ryukyu Islands, coll. Y. Cai and T. Naruse, 15 Jun
2000; 6 males, cl 3.6-3.7 mm, 8 females, cl 4.8-5.5 mm,
3 ovigerous females, cl 5.0-5.3 mm, ZRC 2004.0528,
24817.819N 124833.769E, pH 7.7, Kara Stream, Ishigaki
Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng and T. Naruse,
17 Jun 2000; 1 male, cl 4.3 mm, ZRC 2004.0529, Taiho
River, Okinawa Island, Ryukyu Islands, coll. P. K. L. Ng
et al., 31 May 1998; 2 females, cl 5.0-5.6 mm, 2 ovigerous
females, cl 5.3-6.1 mm, ZRC, Okuma River, Okinawa
Island, Ryukyu Islands, coll. P. K. L. Ng et al., 31 May 1998.
Comparative Material Examined.—Caridina longirostris
H. Milne Edwards, 1837: one male, cl 3.3 mm, 2 females,
syntypes, MNHN Na746, Macta, Near Oran, Algeria, coll.
P. Roux, no date; 2 males, cl 2.3 mm, 12 females, cl 2.8-3.7
mm, 2 ovigerous females, cl 2.7-3.7 mm, syntypes, ZMA
DE 240085, Macta, near Oran, Algeria, coll. P. Roux, no
date. Caridina gracilipes De Man, 1892: 1 ovigerous female, cl 5.3 mm, 2 females, cl 4.2-4.4 mm, syntypes of
Caridina var. gracilipes De Man, 1892a, RMNH D 1317,
Maros River, Sulawesi, Indonesia, coll. M. Weber, Sep.Oct., 1888. 38 specimens, ZMA De 102636, Balingnipa
Brook, Sulawesi, Indonesia, coll. M. Weber, 1888.
Description.—Rostrum long, reaching beyond end of
scaphocerite, anterior 1/3-1/2 slightly upturned, rostral
formula: 1-2þ16-19þ1-2/8-21 (mode 13-19), normally unarmed 1/3 length of rostrum at anterior dorsal margin
between subapical and other teeth; antennal spine distinctly
ventral to inferior orbital angle; pterygostomian angle subrectangular or slightly produced. Sixth pleomore 0.67 times
as long as carapace, 1.6 times as long as fifth somite, slightly
shorter than telson. Telson 4.0 times as long as wide,
terminating in posteromedian projection; with 3 or 4 pairs of
dorsal spinules situated on distal 3/5, 1 pair of dorsolateral
spines near distal end, 3 or 4 pairs of spines on distal margin, lateral pair longer than sublateral pair, median pair
shortest; preanal carina with distinct spine.
Eyes well developed. Antennular peduncle slender, as
long as or slightly shorter than carapace length; proximal
segment half length of antennular peduncle, intermediate
segment 1.5 times as long as third one; stylocerite reaching
0.8 times length of proximal segment. Scaphocerite 4.2 times
as long as wide, inner margin straight, with distolateral spine
distinctly reaching beyond end of antennular peduncle.
Incisor process of mandible ending in irregular teeth,
molar process truncated. Lower lacinia of maxillula broadly
rounded, upper lacinia elongate, with many distinct teeth on
inner margin, palp slender. Upper endites of maxilla subdivided, palp short, scaphognathite tapering posteriorly with
more than 20 long setae at posterior end. Palp of first
maxilliped ending broadly triangular. Second maxilliped
typical of genus. Third maxilliped reaching level of end of
antennular peduncle, with short exopod, reaching only
slightly beyond end of antepenutimate segment, ultimate
segment distinctly shorter than penultimate segment.
Epipods on first 4 pereiopods. First pereiopod reaching
to end of proximal segment of antennular peduncle, merus
399
3.5-3.7 times as long as broad, as long as carpus; carpus
excavated anteriorly, shorter than chela, 2.4-2.5 times as
long as high; chela 2.2-2.4 times as long as broad; fingers
1.1-1.2 times as long as palm. Second pereiopod reaching
end of intermediate segment of antennular peduncle, merus
shorter than carpus, 5.6-5.8 times as long as broad; carpus
distinctly longer than chela, 1.3 times as long as chela, 5.76.3 times as long as high; chela 2.8-2.9 times as long as
broad; fingers 1.3-1.4 times as long as palm. Third
pereiopod long, slender, reaching beyond end of antennular
peduncle by entire dactylus, propodus distinctly shorter than
merus, 13-15 times as long as broad, 5.0-5.5 times as long as
dactylus; dactylus ending in 2 claws; 2.3-2.9 times as long
as wide (spines included), with 3-5 accessory spines
strongly curved inwards. Fifth pereiopod reaching beyond
end of antennular peduncle by entire dactylus, propodus
distinctly longer than merus, 16-17 times as long as broad,
2.9 times as long as dactylus; dactylus 4.1-4.6 times as long
as wide, with 39-54 spinules on flexor margin.
Endopod of male first pleopod extending to 1/4 length
exopod, rounded, subtriangular, 2.2 times as long as broad,
with distinct appendix interna near distal end of endopod.
Appendix masculina of male second pleopod short, reaching
level of half length of endopod, inner and distal surface
densely lined with long spines; appendix interna at basal 2/5
of appendix masculina, extending to distal 1/5 of appendix
masculina. Diaeresis of uropodal exopod with 12-14
spinules.
Egg size 0.41-0.43 3 0.22-0.23 mm in diameter.
Remarks.—Stimpson (1860) briefly described Caridina
grandirostris from the Loo Choo Islands (¼ Okinawa):
‘‘Rostrum carapace vix brevius, appendices antennarum
superans, extremitate gracile paullo reflexum; crista dorsali
supra oculos fere recta et denticulis minutis ad 20 serrata,
denticulo postico supra basim pedunculorum oculorum sito;
cristae parte quarta anteriore edentula, denticulo uno
mediano et duobus apicalibus exceptis; rostri margine
inferiore obscure 8-10 denticulato. Pedum primi paris
carpus quam manus multo brevior; secundi paris carpus
valde gracilis et manu parce longior. Segmentum caudale
lamellis lateralibus quarta parte brevius, dorso paribus sex
aculeorum instructum.’’ [Rostrum rarely shorter than carapace, reaching beyond end of antennular peduncle, anterior
portion slender, slightly upturned; posterior portion of
dorsal margin crested over orbit, armed with about 20 small
teeth, with fewer of them located at carapace posterior to
orbital margin; anterior quarter no teeth, except 1 or 2 apical
teeth and subapical teeth; ventral margin armed with 8-10
very small teeth. Carpus of first pereiopod much shorter than
dactylus; carpus of second pereiopod very slender, longer
than dactylus. Telson 4 times as long as wide, dorsal surface
equipped with 6 spinules.]. Our specimens agree generally
with the original description, although some variation could
be observed: the number of teeth on the ventral margin of
the rostrum is 8 to 21 in our specimens (vs. 8-10 in
Stimpson’s) and the unarmed anterior dorsal margin of the
rostrum measured from 1/4 to 1/2 (vs. 1/4 in Stimpson’s).
These differences can easily be explained by the intraspecific variation among populations and the fact that
Stimpson only had a small sample size from one location.
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CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
With regard to the possession of small eggs, a telson
which terminates in a posteromedian projection and the
preanal carina having a distinct spine, C. grandirostris most
closely resembles C. longirostris H. Milne Edwards, 1837,
and C. gracilipes De Man, 1892. However, it differs from
both species by the endopod of the male first pleopod
possessing a distinct appendix interna. It can also be distinguished from C. longirostris by the more slender carpus
of the first pereiopod (2.4-2.5 times as long as high vs. 1.5
times in C. longirostris); the smaller number of spines on
the dactylus of the third pereiopod (5-7 vs. 7-10), the
presence of more spinules on the dactylus of the fifth
pereiopod (40-55 vs. 35-40 in C. longirostris), and slightly
larger eggs (0.41-0.43 3 0.220.23 mm vs. 0.35-0.38 3
0.21-0.26 mm). Stimpson (1860) did compare C. grandirostris with C. longirostris, commenting that ‘‘A C.
longirostris differt dentibus rostri superne margis numerosis.’’ This difference, however, is not valid. Studies of
a large number of specimens show that the number of spines
on the dorsal margin of the rostrum ranges from 8-21 in C.
grandirostris and 14-21 in C. longirostris. Caridina
grandirostris can further be separated from C. gracilipes
by the more slender carpus of the first pereiopod (2.4-2.5
times as long as high vs. 2.0-2.3 times in C. gracilipes); the
relatively shorter fingers of the first pereiopod (1.1-1.2 times
as long as palm vs. 1.3-1.7 in C. gracilipes); the shorter and
stouter dactylus of the third pereiopod (propodus 4.5-5.0
times as long as dactylus vs. 3.8-4.4 times in C. gracilipes;
dactylus 2.3-2.9 times as long as wide vs, 4.2-4.8 times in C.
gracilipes); the smaller number of spines on the dactylus of
the third pereiopod (5-7 vs. 9-10); and the longer and stouter
dactylus of the fifth pereiopod (propodus 2.9 times as long
as dactylus vs. 3.7-3.8 times in C. gracilipes; dactylus 4.14.6 times as long as wide vs. 5.0-6.0 times in C. gracilipes).
Although Kubo (1938) rediagnosed C. grandirostris, all
subsequent authors have used the name C. nilotica (see
Fujino, 1972; Miyake, 1977; Shokita, 1975) or C. longirostris (see Shokita, 1979; Hayashi, 1989c; Suzuki and
Sato, 1994) instead. Since the type specimens of C.
grandirostris are lost, and because of the taxonomic problems discussed, a specimen from Tima River of Okinawa
Island is herein designated as the neotype of Caridina
grandirostris, to stabilize the taxonomy of the species. The
neotype, a male (cl 4.9 mm), from Tima River of Okinawa
Island, Ryukyu Islands, is deposited in the NSMT.
Distribution.—Known only from the Ryukyu Islands,
Japan.
Habitat.—Freshwater bodies with connections to the sea.
Caridina leucosticta Stimpson, 1860
Figs. 7-10
Caridina leucosticta Stimpson, 1860: 97 [type locality: Simoda, Izu
Peninsula, Honshu, Japan].
401
Caridina leucosticta Bouvier, 1905: 93; Kubo, 1938: 87, figs. 16a-h; 1986:
493, fig. 20-79-3; Kamita, 1961: 64, figs: 28a-d; Fujino, 1972:7, fig.
19; Shokita, 1975: 119; 1979: 202; Miyake, 1977: 39; Hayashi,
1989b: 230; Suzuki et al., 1993: 56; Suzuki and Sato, 1994: 64.
Not Caridina leucosticta: Kamita, 1967: 5, fig. 5.
Material Examined.—Neotype: ovigerous female, cl 7.7
mm, NSMT (NIES-Iz-1), 138856.439E 34841.839N, Inouzawa River, freshwater, lotic, EC 134, Shimoda City, Izu
Peninsula, coll. K. Satake, 10 Aug 2004. Others: 14 males,
cl 3.9-5.6 mm, 7 ovigerous females, cl 6.9-8.5 mm, NIESIz-1, 138856.439E 34841.839N, Inouzawa River, freshwater,
lotic, EC 134, Shimoda City, Izu Peninsula, coll. K. Satake,
10 Aug 2004; 24 males, cl 3.9-5.0 mm, 10 ovigerous females, cl 4.8-7.3 mm, NIES-Iz-2, 138852.589E 34838.729N,
a small stream draining into Aono River, brackish water,
lotic, EC 3810, Minami-Izu, Izu Peninsula, coll. K. Satake,
10 Aug 2004; 3 ovigerous females, cl 7.5-9.0 mm, NIES-Iz3, 138847.679E 34845.219N, Naka River, freshwater, lotic,
EC 139, Matsuzaki-cho, Izu Peninsula, coll. K. Satake, 10
Aug 2004; 1 male, cl 3.9 mm, NSMT, 26836.619N
128807.049E, Arume River, Okinawa Island, Ryukyu
Islands, coll. Y. Cai, N. K. Ng, T. Naruse and S. Islam,
11 Jun 2000. Others: 1 male, cl 4.4 mm, 1 female, cl 6.3
mm, 2 ovigerous females, cl 5.8-5.9 mm, ZRC 2004.0531,
Okuma River, Okinawa Island, Ryukyu Islands, 31 Jun
1998; 14 males, cl 3.5-3.9 mm, 1 female, cl 4.0 mm, 7
ovigerous females, cl 4.5-5.8 mm, ZRC 2004.0530,
26833.489N 128802.489E, freshwater stream draining to
a small patch of mangrove, Oura River, Okinawa Island,
Ryukyu Islands, coll. Y. Cai et al., 11 Jun 2000; 5 males, cl
3.0-4.0 mm, 20 females, cl 4.5-5.2 mm, 19 Jan 1975, Yona
River, Okinawa Island, Ryukyu Islands; 1 male, cl 48.0 mm,
Aritsu River, Kume-jima, coll. A. Kawakami, 8 Dec 1995; 1
male, cl 3.8 mm, 1 ovigerous female, cl 6.5 mm, ZRC,
24816.649N 123852.809E, shallow freshwater stream of Aira
River, Iriomote Island, Ryukyu Islands, coll. Y. Cai, N. K.
Ng and T. Naruse, 14 Jun 2000; 9 males, cl 3.0-4.2 mm, 12
ovigerous females, cl 4.7-6.5 mm, ZRC 2004.0532,
24816.609N 123852.749E, shallow freshwater stream, downstream of Aira River, Iriomote Island, Ryukyu Islands, coll.
Y. Cai, N. K. Ng and T. Naruse, 14 Jun 2000; 7 males, cl
3.1-3.5 mm, 7 ovigerous females, cl 4.4-4.9 mm, ZRC
2004.0533, 24818.399N 123851.299E, downstream from the
headwater of Nakama River, Iriomote Island, Ryukyu
Islands, coll. Y. Cai, N. K. Ng and T. Naruse, 15 Jun
2000; 6 males, cl 4.2-4.7 mm, 8 females, cl 5.8-6.4 mm, 15
ovigerous females, cl 5.8-6.3 mm, ZRC 2004.0534, fast
flowing water, upper part of Omija River, Iriomote Island,
Ryukyu Islands, coll. Y. Cai and T. Naruse, 15 Jun 2000; 3
males, cl 2.4-4.0 mm, 8 females, cl 4.6-6.4 mm, 1 ovigerous
female, cl 5.7 mm, ZRC 2004.0535, upper Hiji fall, Hiji
River, coll. P. K. L. Ng et al., 4 Nov 1987; 3 females, cl 5.05.9 mm, UR1750, no data; 1 ovigerous female, cl 5.7 mm,
UR1221, Kawauchi-gawa River, Amami-ohshima Island,
Ryukyu Islands, coll. M. Takeda, 15 Jul 1988; 1 female, cl
Fig. 5. Caridina grandirostris. A, first pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fifth pereiopod; F, dactylus of
fifth pereiopod; G, male first pleopod; H, male second pleopod. Scales: A-C, E ¼ 1.0 mm; D, F ¼ 0.2 mm; G, H ¼ 0.5 mm. (neotype male, cl 4.9 mm, NSMT,
Tima River, Okinawa, Ryukyu Islands)
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CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
3.0 mm, UR1181, Sisa-gawa River, Matsuura city, Nagasaki
Pref., Kyushu, no date; 1 female, cl 3.0 mm, Kanyu-gawa
River, Amami-ohshima Island, Ryukyu Islands, coll. M.
Takeda, 16 Jul 1988; 4 males, cl 4.4-5.0 mm, 6 ovigerous
females, cl 6.6-6.8 mm, UR1111, Kawauchi-gawa River,
Amami-oshima Island, Ryukyu Islands, coll. M. Takeda,
18 Jul 1988; 1 ovigerous female, cl 6.2 mm, UR1219,
Kawauchi-gawa River, Amami-ohshima Island, Ryukyu
Islands, 18 Jul 1988.
Comparative Material Examined.—Caridina wyckii (Hickson, 1888): 11 females, cl 3.6-4.1 mm, 1 ovigerous female, cl
3.6 mm, syntypes, RMNH D 1456, Lake Tondano, leg. (S.
Hickson, Apr 1886); Caridina brachydactyla De Man, 1908:
1 female, cl 6.7 mm, 2 ovigerous females, cl 5.1-5.3 mm,
RMNH D 2552, syntype of Caridina nilotica var. brachydactyla, Luwu, River near Palopo, Celebes (Sulawesi), coll.
M. Weber, Feb 1889; Caridina nilotica meridionalis J.
Roux, 1926: 14 males, cl 2.4-5.3 mm, 6 females, cl 2.4-5.7
mm, 1 ovigerous female, cl 5.7 mm, NHMB, syntypes,
Tiouaka Tal, New Caledonia, 20 Aug 1911.
Description.—Rostrum long, reaching to or slightly beyond
end of scaphocerite, straight or anterior 1/3 slightly
upturned, rostral formula: 2-4 (mode 3) þ 15-22 (mode 1518) þ 1-3 (mode 1)/7-13 (mode 8-10), normally unarmed anterior dorsal margin between subapical teeth and others 1/4
length of rostrum or less; antennal spine distinctly lower than
inferior orbital angle; pterygostomian margin produced.
Sixth pleomore 0.8 times as long as carapace, 1.9 times as
long as fifth somite, slightly shorter than telson. Telson 3.2
times as long as wide, terminating in posteromedian projection; with 3 or 4 pairs of dorsal spinules, 1 pair of
dorsolateral spines near distal end, 3 or 4 pairs of spines on
distal margin, lateral pair longer than sublateral pair, median
pair shortest; preanal carina moderately high, without spine.
Eyes well developed. Antennular peduncle slender, 0.7 to
0.9 times as long as carapace; proximal segment a little more
than half length of the antennular peduncle, intermediate
segment 1.6 times as long as distal segment; stylocerite
reaching 0.85 times length of proximal segment. Scaphocerite 3.4 times as long as wide, with distolateral spine
reaching slightly beyond end of antennular peduncle.
Mouthparts similar to C. grandirostris. Third maxilliped
reaching to end of antennular peduncle, with a short exopod,
reaching only slightly beyond end of antepenutimate
segment, ultimate segment distinctly shorter than penultimate segment.
Epipods on first 4 pereiopods. First pereiopod short,
reaching to distal end of eye stalk, merus 2.8-3.2 times as
long as broad, as long as carpus; carpus excavated anteriorly, shorter than chela, 1.9-2.2 times as long as high;
chela 2.0-2.4 times as long as broad; fingers 1.1-1.4 times as
long as palm. Second pereiopod reaching end of second
403
segment of antennular peduncle, merus shorter than carpus,
5.0-5.6 times as long as broad; carpus distinctly longer than
chela, 1.2-1.3 times as long as chela, 4.5-5.5 times as long as
high; chela 2.3-2.8 times as long as broad; fingers 1.2-1.6
times as long as palm. Third pereiopod slender, reaching to
end of antennular peduncle, propodus distinctly shorter than
merus, 13-15 times as long as broad, 4.0-5.2 times as long as
dactylus; dactylus ending in 2 claws; 2.9-3.2 times as long
as wide (spines included), with 5 accessory spines strongly
curved inwards; fifth pereiopod reaching end of antennular
peduncle, propodus distinctly longer than merus, 16-19
times as long as broad, 2.9 times as long as dactylus;
dactylus 3.8-4.8 times as long as wide, with 41-61 spinules
on its flexor margin.
Endopod of male first pleopod reaching 1/4 length of
exopod, subtriangular, 2.0-2.2 times as long as broad, no
appendix interna, or with a vestige. Appendix masculina
of male second pleopod short, reaching to half length of
endopod, inner and distal surfaces densely lined with long
spines; appendix interna at basal 2/5 of appendix masculina,
extending to distal 1/5 of appendix masculina. Diaeresis of
uropodal exopod with 14 or 15 spinules.
Egg size 0.38-0.40 3 0.23-0.28 mm in diameter.
Remarks.—Stimpson (1860) briefly described Caridina
leucosticta from Simoda, Japan, as follows: ‘‘Rostrum
circiter carapacis longitudine, pedunculo antennularum
longius; margine superiore recto, dentibus tenuibus ad
17 þ 3 armato, apicem versus parce resimo et edentulo;
margine inferiore 10 dentato. Spina antennalis alte posia.
Pedes gracillimi; posticorum merus margine inferiore
spinulis 2-5 armatus. Color obscure-fuscus, maculis vel
stigmis minutis crebris albis notatus.’’ [Rostrum as long as
carapace, longer than antennular peduncle; upper margin
straight, armed with 17 small teeth, apical region slightly
upturned, with 3 apical teeth, no teeth between apical and
straight upper margin; ventral margin with 10 teeth.
Antennular spine not located at inferior angle. Legs slender,
inferior margin of merus armed with 2-5 spines, Color
obscure dark, covered with numerous tiny white spots.]. Our
specimens agree well with the original description. With
regard to the small egg size, the form of the rostrum, the lack
of a preanal spine and having a posteromedian projection on
the distal margin of the telson, C. leucosticta appears closest
to C. brachydactyla De Man, 1908. It can be distinguished
from C. brachydactyla by the relatively shorter fingers of
first pereiopod (fingers 1.1-1.4 times as long as palm vs. 2.02.5 times in C. brachydactyla); the lack of a prominent spine
at the distal end of propodus of fifth pereiopod (vs. having
such spine which reaches beyond half of dactylus in C.
brachydactyla); the longer dactylus of third pereiopod
(propodus 4.0-5.2 times as long as dactylus vs. 6.0 times
in C. brachydactyla); and the longer and more slender
dactylus of the fifth pereiopod (propodus 3.8-4.8 times as
Fig. 6. Caridina grandirostris. A, cephalothorax and cephalic appendages, lateral view; B, distal portion of telson; C, preanal carina; 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. Scales: A,
D-F, H ¼ 1.0 mm; C ¼ 0.5 mm; B, G, I, J ¼ 0.2 mm. (ovigerous female, 6.1 mm, ZRC, Okuma River, Okinawa Island, Ryukyu Islands)
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 26, NO. 3, 2006
Fig. 7. Caridina leucosticta. A, cephalothorax and cephalic appendages, lateral view; B, preanal carina; C, distal portion of telson, D, scaphocerite;
E, mandible; F, maxillula; G, maxilla; H, first maxilliped; I, second maxilliped; J, third maxilliped; K, uropodal diaeresis. Scales: A, D ¼ 1.0 mm; B, E-J ¼
0.5 mm, C, K ¼ 0.2 mm. (neotype: ovigerous female, cl 7.7 mm, NSMT, Inouzawa River, freshwater, Shimoda, Izu Peninsula)
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
long as dactylus vs. 5.0-6.0 times in C. brachydactyla;
dactylus 3.8-4.8 times as long as wide vs. 3.4-3.8 times
in C. brachydactyla). Bouvier (1905) commented that with
respect to the straight rostrum, C. leucosticta strongly
resembles C. wyckii (Hickson, 1888). However, C. leucosticta can be immediately distinguished from C. wyckii by
its longer rostrum which reaches or is slightly beyond the
end of scaphocerite (vs. not reaching beyond end of
antennular peduncle); the stouter scaphocerite (3.4 times
as long as wide vs. 4.3 times); the more slender carpus of the
first pereiopod (1.9-2.2 times as long as high vs. 1.4-1.5);
the preanal carina without a spine (vs. with spine), and the
relatively smaller egg size (0.38-0.40 vs. 0.23-0.28 mm vs.
0.52 3 0.30 mm). Caridina leucosticta also resembles C.
nilotica meridionalis Roux, 1926, from New Caledonia, in
the form of the rostrum, but differs from the latter by lacking
an appendix interna on the male first pleopod and having a
smaller egg size (0.38-0.40 3 0.23-0.28 mm vs. 0.65-0.70 3
0.40 mm).
Kamita (1967) reported C. leucosticta from New
Caledonia on the basis of one female specimen. This record
was subsequently cited by Holthuis (1969). Kamita’s (1967)
specimen has a rostrum, which is shorter than the antennular
peduncle, with 15 ventral teeth, while in C. leucosticta, the
ventral teeth are always no more than 13. Kamita’s (1967)
material is more likely to be C. nilotica meridionalis rather
than C. leucosticta.
In view of the potential for taxonomic problems, an
ovigerous female, cl 7.7 mm, collected from Inouzawa
River, Shimoda City, Izu Peninsula, is herein designated as
the neotype of C. leucosticta Stimpson, 1860. It is deposited
in National Science Museum, Tokyo, Japan.
Distribution.—Known with certainty only from Japan.
Reported from Korea.
Habitat.—Caridina leucosticta lives in rivers which are
connected to the sea, in freshwater or brackish waters. When
collected together with C. grandirostris from the same river
in the Ryukyus, C. leucosticta is usually present in the upper
regions, while C. grandirostris occupies the lower parts. In
some cases, both were caught together. The similar
distribution pattern is also found in Izu Peninsula. However,
the species collected together with it is Paratya compressa
De Haan, 1849.
Caridina multidentata Stimpson, 1860
Figs. 11, 12
Caridina multidentata Stimpson, 1860: 98 [type locality: Ogasawara
(Bonin) Islands, Japan].
Caridina multidentata: Chace, 1997: 16 (part).
Caridina japonica De Man, 1892b: 261, pl. IX, figs. 7, 8 [type locality:
Kagar, Hayagana, Japan]; Bouvier, 1904: 133; 1905: 75; 1925: 239,
figs. 550-554; Kubo, 1938: 89, figs. 17, 18a-h; Kamita, 1956: 14;
1959: 21, figs. 1-3; 1961: 50, figs. 20-27; Holthuis, 1965: 12, fig. 4;
Suzuki, 1972: 6, figs. 4-7; Fujino, 1972: 7, fig. 11; Shokita, 1975:
119; 1979: 203, pl.1g; Hayashi and Hamano, 1984: 571; Hayashi,
1989b: 229, figs. 166a-h; Choy, 1991: 353; Hung et al., 1993: 495,
fig 9a, 10a-f; Suzuki, et al., 1993: 58; Suzuki and Sato, 1994: 68;
Shy and Yu, 1998: 55, fig 23.
Caridina japonica sikokuensis Kubo, 1938: 91, figs. 20a-m [Ryugado,
Kochi Prefecture, Shikoku, Japan].
Not Caridina multidentata De Man, 1892a: 380, pl. 22, fig. 26; Bouvier,
1905: 74.
405
Material Examined.—Neotype of Caridina multidentata:
ovigerous female, cl 9.8 mm, NSMT, from a stream in
Tenno-ura, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake and H. Suzuki, 4 May 2000. 1 male, cl 8.0 mm,
lectotype of Caridina japonica De Man, 1892b, ZMA De
102876, Japan, coll. J. Anderson, 1881, 4 males, cl 7.8-8.2
mm; paralectotype of Caridina japonica De Man, 1892b,
data same as lectotype; 1 male, cl 8.8 mm, paralectotype,
MNHN-Na 731, Kagar, Hayagana, Japan. Others: 4 males,
cl 6.0-7.4 mm, 16 ovigerous females, cl 7.5-9.1 mm, NIESH-3, Nagahama-bashi, lotic system, EC 270-610, 130 to 140
meters above sea level, Haha-jima Island, Ogasawara Islands, coll. K. Satake and R. Ueno, 2 Mar 1999; 23
ovigerous females, cl 6.5-9.0 mm, NIES-H-6, upper reach of
Koromodate Stream, lotic system, EC 590-700, 70 to 110
meters above sea level, Haha-jima Island, Ogasawara
Islands, coll. K. Satake and R. Ueno, 24 Feb 2000; 1 male,
cl 7.4 mm, 1 ovigerous female, cl 7.9 mm, NIES-H-9, an
inlet stream of Chibusa Dam, lotic system, EC 250-600; 60
meters above sea level, Haha-jima Island, Ogasawara
Islands, coll. K. Satake, 25 Feb 2000; 4 males, cl 6.3-7.5
mm, NIES-H-10, an inlet stream of Chibusa East Dam, lotic
system, EC290-550, 70 meters above sea level, Haha-jima
Island, Ogasawara Islands, coll. K. Satake and R. Ueno, 26
Feb 2000; 3 males, cl 7.4-8.0 mm, NIES-CH-12, east inlet
stream of Shigure Dam, lotic, EC 200, 90 meters above sea
level, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake, R. Ueno, K. Horikoshi and M. Inaba, 27 Feb
2000; 5 females, cl 6.8-9.5 mm, 8 males, cl 6.2-8.0 mm,
NIES-CH-5, Minamifukuro-zawa stream, lotic system, EC
960-1200, 0 to 10 meters above sea level, Chichi-jima
Island, Ogasawara Islands, coll. K. Satake and R. Ueno, 11
Feb 1998; 14 males, cl 5.1-6.0 mm, 10 females, cl 5.9-9.0
mm, NIES-CH-17, one stream of Naka-kaigan Coast,
freshwater, lotic system, EC 380, 60 to 70 meters above
sea level, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake, R. Ueno and M. Inaba, 28 Feb 2000; 20 males, cl
5.8-8.5 mm, 8 females, cl 7.3-9.1 mm, 12 ovigerous
females, cl 7.8-9.5 mm, NIES-CH-18, a stream of Higashikaigan Coast, lotic system, EC 400-510, 140 meters above
sea level, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake, 29 Feb 2000; 7 males, cl 6.1-6.9 mm, 2 females, cl
6.6-9.8 mm, NIES-CH-19, one tributary of a stream of
Higashi-kaigan Coast, lotic system, EC 420, 250 meters
above sea level, Chichi-jima Island, Ogasawara Islands,
coll. K. Satake and M. Inaba, 28 Apr 2000; 1 female, cl 7.2
mm, NIES-CH-20, the other tributary of a stream of
Higashi-kaigan Coast, lotic, EC 280, 200 meters above the
sea level Chichi-jima Island, Ogasawara Islands, coll. K.
Satake and M. Inaba, 3 May 2000; 26 males, cl 5.7-6.5 mm,
1 females, 3.2-8.6 mm, NIES-CH-23, a stream in Tenno-ura,
lotic system, EC 610, 90 meters above sea level, Chichijima Island, Ogasawara Islands, coll. K. Satake and H.
Suzuki, 4 May 2000; 1 male, cl 5.7 mm, 3 ovigerous
females, cl 7.0-8.4 mm, ZRC 2004.0536, from a stream in
Tenno-ura, Chichi-jima Island, Ogasawara Islands, coll. K.
Satake and H. Suzuki, 4 May 2000; 1 ovigerous female, cl
11.0 mm, NSMT-Cr1673, 338069150N 1398489170E, Kamogawa River, Hachijo-jima Island, Izu Archipelago, coll. J.
Okuno, 15 July 1992; 7 males, cl 7.2-8.2 mm, NSMT-
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 26, NO. 3, 2006
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
Cr2144, 338069150N 1398489170E, Kamo-gawa River,
Hachijo-jima Island, Izu Islands, coll. J. Okuno, 25 Sep
92; 4 ovigerous females, cl 9.5-11.5 mm, NSMT-Cr1674,
338069150N 1398489170E, Kamo-gawa River, Hachijo-jima
Island, Izu Island, coll. J. Okuno, 15 Sep 1992; 1 male, cl
11.3 mm, NSMT-Cr1542, upper stream of Kochi-gawa
River, Numazu, Izu Peninsula, Honshu, 200 meters above
sea level, coll. N. Seno, 2 Nov 1987; 1 female, cl 9.9 mm,
NSMT-Cr1745, no data; 1 male, cl 7.5 mm, NSMT-Cr1121,
upper of Kawauchi-gawa River, Amami-oshima Island,
Ryukyu Islands, coll. M. Takeda, 15 Jul 88; 1 ovigerous
female, cl 7.9 mm, NSMT-Cr1754, Shino-Kawa River,
Amami-oshima Island, Ryukyu Islands, coll. M. Takeda, 17
Jul 1988; 1 male, cl 8.3 mm, 2 ovigerous females, cl 7.3-9.8
mm, ZRC 2004.0538, Omija River, Iriomote Island,
Ryukyu Islands, coll. T. Naruse, 1 Aug 1999; 1 ovigerous
female, cl 8.2 mm, NSMT, Amami-ohshima Island, Ryukyu
Islands, 29 Jul 1992; 1 male, cl 16 mm, 2 ovigerous females,
cl 10-12 mm, ZRC, 26839.949N 128815.319E, Arakawa
River, Okinawa Island, Ryukyu Islands, coll. Y. Cai, N. K.
Ng, T. Naruse and S. Islam, 11 Jun 2000; 1 female, cl 4.9
mm, ZRC, fast flowing water, upper part of Omija
River, Iriomote Island, Ryukyu Islands, coll. Y. Cai and
T. Naruse, 15 Jun 2000; 1 female, cl 9.5 mm, 2 ovigerous
females, 6.2-9.8 mm, ZRC 2004.0539, Taiho River,
Okinawa Island, Ryukyu Islands, coll. P. K. L. Ng et al.,
31 May 1998.
Description.—Rostrum reaching slightly beyond end of
basal segment of antennular peduncle to near end of third
segment at most, posterior 2/3 sloping down, with a crest
over orbit, anterior 1/3 horizontal; armed normally throughout both margins except subapical region, rostral formula:
15-30/5-18; antennal spine fused with inferior orbital angle;
pterygostomian margin subrectangular. Sixth abdominal
somite 0.60 times as long as carapace, 1.5 times as long as
fifth somite, slightly shorter than telson. Telson 3.2 times as
long as wide, terminating in a posteromedian projection; 3
or 4 pairs of dorsal spinules, situated on posterior half, 1 pair
of dorsolateral spines near distal end, 3 or 4 pairs of spines
on distal margin, lateral pair subequal to intermediate pairs;
preanal carina high, without spine.
Eyes well developed. Antennular peduncle slender, 0.8
times as long as carapace; basal segment a little more than
half length of the antennular peduncle, second segment 1.5
times as long as third segment; stylocerite reaching 0.8 times
length of basal segment. Scaphocerite 2.7 times as long as
wide, with distolateral spine reaching end of antennular
peduncle.
Third maxilliped reaching to end of third segment of
antennular peduncle, with a short exopod, reaching only
slightly beyond end of antepenutimate segment, ultimate
segment distinctly shorter than penultimate segment.
Epipods on first 4 pereiopods. First pereiopod short,
reaching to end of basal segment of antennular peduncle,
407
merus 2.0-2.2 as long as broad, slightly longer than carpus;
carpus excavated strongly anteriorly, shorter than chela, 1.11.3 times as long as high; chela 2.0-2.3 times as long as
broad; length of fingers variable, at most, shorter than palm,
sometimes as long as or even slightly longer. Second
pereiopod reaching end of second segment of antennular
peduncle, merus shorter than carpus, 3.9-4.4 times as long
as broad; carpus slightly longer than chela, 3.8-4.0 times as
long as high; chela 2.5-2.7 times as long as broad; fingers
1.5-1.7 times as long as palm. Third pereiopod long,
slender, reaching beyond end of antennular peduncle by
entire dactylus and 1/4 of propodus, propodus distinctly
shorter than merus, 10-15 times as long as broad, 3.7-4.4
times as long as dactylus; dactylus ending in a strong claw;
2.0-2.4 times as long as wide (spines included), flexor
margin with 5 or 6 accessory spines strongly curved inwards, distal accessory spine distinctly shorter than the
following one. Fifth pereiopod reaching to end of antennular
peduncle, propodus distinctly longer than merus, 11-15
times as long as broad, 5.3 times as long-as-dactylus; distal
end of propodus normally with an enlarged spine reaching
to half or 3/4 of the dactylus length; dactylus 2.2-2.6 times
as long as wide, with 35-48 spinules on flexor margin.
Endopod of male first pleopod extending to 1/3 length of
exopod, rounded, subrectangular, 2.0-2.3 times as long as
broad, with a distinct appendix interna near distal end of
endopod. Appendix masculina of male second pleopod
short, reaching to half length of endopod, inner and distal
surface densely lined with long spines; appendix interna
at basal 1/3 of appendix masculina, extending to distal 1/3
of appendix masculina. Diaeresis of uropodal exopod with
19-23 spinules.
Egg size 0.38-0.40 3 0.23-0.28 mm in diameter.
Remarks.—Stimpson (1860) described C. multidentata from
Bonin Islands (Ogasawara Islands) in the following words:
‘‘Rostrum medium articuli ultimi pedunculi antennularum
attingens; crista dorsai lamellato-dilatata, arcuata, supra
bases oculorum oriente, et denticulis 20-30 serrata;
extremitate robusta, acuta, vix denticulata; margin inferior
14-denticulato. Margo carapacis anterior spina antennali
armatus. Pedes secundi paris pedunculum antennularum
superantes; carpo manu longiore; digitis degressis, penicillis
densis, latis, fere flabelliformibus. Dactyli pedum posticorum breves, septimam partem articuli penultimi longitudine
non superantes. Segmentum caudale dorso non concavum,
paribus quinque aculeorum instructum; lamellae laterales
grandes, segmento caudale fere duplo longiores, extremitatibus productis subtriangularibus.’’ [Rostrum medium,
reaching to utimate segment of antennular peduncle, dorsal
crest ridged dilated, bending against base of ocular region,
serrated with 20-30 denticles; upper surface stout, sharp,
heavily denticulate; lower margin with 14 denticles. Margin
of carapace armed anteriorly with antennal spine. Second
pair of legs longer than antennular peduncle; carpus elon-
Fig. 8. Caridina leucosticta. A, first pereiopod; B, second pereiopod; C, third pereiopod; D, dactylus of third pereiopod; E, fifth pereiopod; F, dactylus of
fifth pereiopod; G, male first pleopod; H, male second pleopod. Scales: A, B ¼ 0.5 mm, C, E ¼ 1.0 mm; D, F ¼ 0.2 mm; G, H ¼ 0.2 mm. (neotype: ovigerous
female, cl 7.7 mm, NSMT, Inouzawa River, freshwater, Shimoda, Izu Peninsula).
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Fig. 9. Caridina leucosticta. 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 masculina; J, preanal carina; K, distal portion of telson. Scales: A-D, F ¼ 1.0 mm; H, I ¼ 0.5 mm; E, G, K ¼ 0.2 mm. (male, cl 4.0 mm,
ZRC 2004.0535, Hiji River, Okinawa Island, Ryukyu Islands)
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
409
Fig. 10. Caridina leucosticta. 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, cephalothorax and cephalic appendages, lateral view; B, telson C, distal
portion of telson; D, mandible; E, maxillula; F, maxilla; G, first maxilliped; H, cephalothorax and cephalic appendages, lateral view; I, first pereiopod;
J, second pereiopod; K, dactylus of third pereiopod. Scales: A-C, D, F, H-J ¼ 1.0 mm; E, G, K ¼ 0.2 mm. (A-G, female, cl 5.2 mm, ZRC 2004.0535, Hiji
River, Okinawa Island, Ryukyu Islands; H-K, female, cl 6.1 mm, ZRC 2004.0535, Hiji River, Okinawa Island, Ryukyu Islands)
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Fig. 11. Caridina multidentata. A, F, cephalothorax and cephalic appendages, lateral view; B, telson; C, distal portion of telson; D, male first pleopod;
E, male second pleopod. A, B, F ¼ 1.0 mm; D, E ¼ 0.5 mm; C ¼ 0.2 mm. (A-E, male, cl 5.7 mm, F, ovigerous female, cl 9.8 mm, neotype, NSMT, from a
stream of Tenno-ura, Chichi-jima Island, Ogasawara Islands)
!
Fig. 12. Caridina multidentata. A, first pereiopod; B, second pereiopod; C, fifth pereiopod; D, dactylus of fifth pereiopod; E, third pereiopod; F, dactylus
of third pereiopod; H, uropodal diaeresis. Scales: A-C, G ¼ 0.5 mm; E ¼ 1.0 mm; D, F, H ¼ 0.2 mm (male, cl 5.7 mm, ZRC 2004.0536, from a stream of
Tenno-ura, Chichi-jima Island, Ogasawara Islands)
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gate, fingers small, thick pointed, flattened, tip expanded.
Dactylus of posterior legs sharp, penultimate article of
seventh part longitudinally not produced. Caudal segment
dorsally not concave, equal in size to fifth, with large lateral
ridges, tip of caudal segment twice as long, triangular.]
Surprisingly, the name C. multidentata Stimpson, 1860,
has been totally ignored in treatments of the crustacean fauna
of Japan since its original report (e.g., see Kubo, 1938;
Kamita, 1956; Shokita, 1975; Suzuki, 1972; Hayashi, 1989ac, 1990). De Man (1892a), however, reported C. multidentata from Indonesia, on which Chace (1997) stated that
its identification must be considered questionable for the
time being. According to the description and figures (De
Man, 1892a; Bouvier, 1925), the Indonesian material
distinctively differs from C. multidentata in having a
straight, not crested rostrum (vs. crested), smaller number
of uropodal teeth on the diaeresis (13-14 vs. 19-23), and the
larger egg size (near 1 mm in diameter vs. 0.38-0.40 3 0.230.28 mm). De Man’s material appears to belong to an
undescribed taxon.
De Man (1892b) described C. japonica on the basis of six
specimens from Kagar, Hayagana, Japan. According to his
description, the upper margin of the rostrum is straight or
very slightly concave, with a rostral formula of 15-22/5-18,
and the pereiopods are of the stout form. In fact, this species
is almost identical to C. multidentata as originally described
by Stimpson (1860). The re-examination of the extant
syntypes in ZMA and MNHN confirms this synonymization. Thus, the specimens previously identified under the
name C. japonica should be reassigned to C. multidentata
instead. Holthuis (1965) stated that a ‘‘. . . comparison of the
Madagascar material with the good description and figures
given by Kubo (1938: 89, figs. 17-19) of Japanese
specimens failed to produce any difference that might be
considered of specific or subspecific value.’’ When Hung
et al. (1993) reported C. japonica from Taiwan, they
commented that it is distinct in having a basal crest on the
rostrum. However, the material from Madagascar reported
by Holthuis (1965: fig-4a) lacks such a crest. A comparison
of the Indian Ocean material with that from the Western
Pacific will be necessary to ascertain whether the present
species truely has such a wide geographical distribution.
Two females of Holthuis’ (1965) material were reexamined. One has a straight rostrum, while the other one
has a small crest over the eyes, a characteristic dactylus on
the third pereiopod and there is an enlarged spine on the
large specimen. All these clearly indicate that the Madagascar specimens are in fact C. multidentata (¼ C. japonica)
as presently defined. Kubo (1938) described a new subspecies, C. japonica sikokuensis, on the basis of specimens
living in a limestone cave, Ryugado, Shikoku. However, the
only characters used to separate this subspecies; the shorter
and sharply bent rostrum, fall easily within the known
variation for C. multidentata as observed here. Kamita
(1956) commented that the typical form could also be
collected in the waters inside and outside of the cave.
Therefore it is likely that this subspecies is merely a variation
form of the typical species. Choy (1991: 354) reported the
occurrence of C. japonica in Fiji and commented that ‘‘A
comparison of original descriptions with the Fijian material
indicate that the morphometric and meristic variations of
many of the characters are large and overlap considerably
thus not justifying the separation of the Fijian form to
a distinct subspecies’’, although he found that there are some
minor differences in the dactylus of the last three pereiopods
and in the placement of rostral teeth.
To stabilize the taxonomy of this species, an ovigerous
female, with a carapace length of 8.8 mm (NSMT), from
a stream in Tenno-ura, Chichi-jima Island, Ogasawara
Islands, is designated as the neotype of Caridina multidentata Stimpson, 1860. A lectotype for Caridina japonica
is also selected (male, cl 8.0 mm, ZMA).
Distribution.—Main islands of Japan, Ogasawara Islands,
Ryukyu Islands, Taiwan, Fiji and Madagascar.
Habitat.—Caridina multidentata lives in large rivers with
big rocks on the substrate.
Caridina typus H. Milne Edwards, 1837
Figs. 13-15
Caridina typus H. Milne Edwards, 1837: 363, Pl. 25, figs. 4, 5 [type
locality: unknown].
Caridina typus: Bouvier, 1904: 133; Kamita, 1959: 21; Ooishi, 1970: 87;
Suzuki, 1972: 10, figs. 8-10; Fujino, 1972: 7, fig. 8; Shokita, 1975:
119; 1979: 204, pl. 1e; Hayashi, 1989c: 310, fig. 168; Suzuki et al.,
1993: 56; Suzuki and Sato, 1994: 60.
Caridina typa: Bouvier, 1905: 77.
Caridina exilirostris Stimpson, 1860: 98 [type locality: Okinawa (Loo
Choo) Island, Ryukyu Islands, Japan]; Bouvier, 1904: 134.
Material Examined.—Syntypes of Caridina typus H. Milne
Edwards: 3 females, cl 7.0-8.4 males, MNHN-Na 930,
locality unknown, no date. Neotype of Caridina exilirostris:
ovigerous female, cl 7.4 mm, NSMT, Okuma River,
Okinawa Island, Ryukyu Islands, coll. P. K. L. Ng et al.,
31 May 1998. Others: 28 males, cl 3.3-5.1 mm, 2 females, cl
5.7-5.8 mm, 10 ovigerous females, cl 6.5-8.3 mm, 10 males,
cl 3.7-6.6 mm, NIES-H-6, upper reach of Koromodate
Stream, lotic system, 70 to 110 meters above sea level,
Haha-jima Island, Ogasawara Islands, coll. K. Satake and R.
Ueno, 24 Feb 2000; 1 male, cl 5.0 mm, 1 female, cl 6.0 mm,
NIES-H-10, an inlet stream of Chibusa East Dam, lotic
system, 70 meters above sea level, Haha-jima Island,
Ogasawara Islands, coll. K. Satake and R. Ueno, 26 Feb
2000; 2 females, cl 7.0-8.1 mm, 1 ovigerous female 7.0 mm,
NIES-H-11, one tributary of a stream of Iguma Bay, 310
meters above sea level, Haha-jima Island, Ogasawara
Islands, coll. K. Satake, 30 Apr 2000; 3 males, cl 4.7-5.3
mm, 1 female, cl 7.6 mm, NIES-H-119, another tributary of
!
Fig. 13. Caridina typus. A, cephalothorax and cephalic appendages, lateral view; B, preanal carina; C, scaphocerite; 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 of male second pleopod. Scales: A, D-F, H ¼ 1.0 mm; B, C ¼ 0.5 mm; G, I-K ¼ 0.2 mm. (male, cl 5.3 mm, ZRC 2004.0560, Okuma River,
Okinawa island, Ryukyu Islands)
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Fig. 14. Caridina typus. 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, uropodal diaeresis. Scales: A ¼ 0.5 mm; B, F, H, I ¼
0.2 mm; C-E, G ¼ 1.0 mm. (ovigerous female, cl 7.4 mm, neotype of Caridina exilirostris, NSMT, Okuma River, Okinawa island, Ryukyu Islands)
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
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Fig. 15. Caridina typus. A, cephalothorax and cephalic appendages, lateral view; B, preanal carina; C, telson; D, distal portion of telson; E, first pereiopod;
F, second pereiopod; G, third pereiopod; H, dactylus of third pereiopod; I, fifth pereiopod; J, dactylus of fifth pereiopod. Scales: A, C, E-G, I ¼ 1.0 mm; B ¼
0.5 mm, D, H, J ¼ 0.2 mm. (male, cl 7.0 mm, NIES-H-11, a stream to Iguma Bay, Haha-jima Island, Ogasawara Islands)
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the stream of Iguma Bay, 270 meters above sea level, Hahajima Island, Ogasawara Islands, coll. K. Satake, 30 Apr
2000; 43 males, cl 3.3-5.5 mm, 11 females, cl 6.7-8.3 mm,
5 ovigerous females, cl 7.2-9.8 mm, NIES-H-12, an inlet
stream of Oh-zawa Dam, lotic, EC 230, 230 to 240 meters
above sea level, Haha-jima Island, Ogasawara Islands, coll.
K. Satake, 2 May 2000; 1 male, cl 4.8 mm, NIES-CH-2,
Oku-ohtaki, a tributary of west inlet stream of Shigure Dam,
lotic system, 210 to 220 meters above sea level, Chichi-jima
Island, Ogasawara Islands, coll. K. Satake and M. Inaba,
3 May 2000; 4 males, cl 6.7-6.9 mm, NIES-CH-26, upper
reach of a stream of Higashi-kaigan Coast, lotic, 200 metres
above sea level, Chichi-jima Island, Ogasawara Islands,
coll. K. Satake and M. Inaba, 3 May 2000; 34 males, cl 3.66.8 mm, NIES-CH-17, one stream of Naka-kaigan Coast,
lotic system, EC 380, 60 to 70 meters above sea level,
Chichi-jima Island, Ogasawara Islands, coll. K. Satake, R.
Ueno and M. Inaba, 28 Feb 2000; 6 males, 4.0-4.6 mm, 1
female, cl 6.1 mm; 1 ovigerous female, cl 7.1 mm, NIESCH-19, one tributary of a stream of Higashi-kaigan Coast,
250 meters above the sea level, Chichi-jima Island,
Ogasawara Islands, coll. K. Satake and M. Inaba, 28 Apr
2000; 6 males, cl 2.2-4.5 mm, NIES-CH-23, a stream of
Tenno-ura, lotic system, 90 meters above sea level, Chichijima Island, Ogasawara Islands, coll. K. Satake and H.
Suzuki, 4 May 2000; 34 males, cl 3.2-5.8 mm, 1 female, cl
4.7 mm, NIES-CH-20, tributary of a stream of Higashikaigan Coast, lotic system, 200 meters above sea level,
Chichi-jima Island, Ogasawara Islands, coll. K. Satake and
M. Inaba, 3 May 2000; 10 males, cl 3.0-4.1 mm, 2 females,
cl 4.7-5.0 mm, NIES-CH-21, stream at Naka-kaigan Coast,
150 meters above the sea level, Chichi-jima Island,
Ogasawara Islands, coll. K. Satake and H. Suzuki, 4 May
2000; 27 males, 3.7-8.1 mm, NIES-CH-13, upper reach of
Tokoyo Fall, 90 to 110 meters above sea level, Chichi-jima
Island, Ogasawara Islands, coll. K. Satake, R. Ueno, K.
Horikoshi and M. Inaba, 27 Feb 2000; 4 males, cl 5.5-7.0
mm, NIES-CH-15, Fukiage-dani Dam, lotic system, 50 to
60 meters above sea level, Chichi-jima Island, Ogasawara
Islands, coll. K. Satake, R. Ueno and Inaba, 23 Feb 2000; 21
ovigerous females, cl 5.6-6.9 mm, 9 females, cl 4.0-6.3 mm,
50 males, cl 3.8-5.0 mm, ZRC 2004.0540, 26833.429N
128804.609E, upstream of Tima River, about 1-2 km from
river mouth, Okinawa Island, Ryukyu Islands, pH 7.4, coll.
Y. Cai, N. K. Ng, T. Naruse and S. Islam, 11 Jun 2000; 3
males, cl 3.7-4.0 mm, ZRC 2004.0541, 26833.639N
128805.529E, upstream of Tima River, about 3 km from
the river mouth, Okinawa Island, Ryukyu Islands, pH 7.4,
coll. N. K. Ng and S. Islam, 11 Jun 2000; 83 males, cl 4.35.7 mm, 1 female, 18 ovigerous females, cl 6.3-7.1 mm,
ZRC 2004.0542, upper stream of Tima River, about 4 km
from river mouth, Okinawa Island, Ryukyu Islands, coll. Y.
Cai, N. K. Ng, T. Naruse and S. Islam, 11 Jun 2000; 42
males, cl 3.8-5.2 mm, 3 females, cl 5.7-6.3 mm, 5 ovigerous
females, cl 6.4-7.0 mm, ZRC 2004.0543, 26839.949N
128815.319E, Arakawa River, Okinawa Island, Ryukyu
Islands, coll. Y. Cai, N. K. Ng, T. Naruse and S. Islam, 11
Jun 2000; 1 male, cl 4.7 mm, 1 female, cl 4.6 mm, 3
ovigerous females, cl 5.2-7.2 mm, ZRC 2004.0544,
24830.119N 124815.269E, Gaburumata River, Ishigaki
Island, Ryukyu Islands, pH 7.3, coll. Y. Cai and T. Naruse,
13 Jun 2000; 1 male, cl 4.5 mm, 1 female, cl 7.0 mm, 4
juveniles, ZRC 2004.0545, 24824.429N 124809.809E, fast
flowing stream, with pH 7.6 at one of the tributaries of
Nagura River, below the reservoir, Ishigaki Island, Ryukyu
Islands, coll. Y. Cai and T. Naruse, 13 Jun 2000; 50 males,
cl 3.5-5.7 mm, 29 females, cl 4.5-7.5 mm, 10 ovigerous
females, cl 5.6-7.9 mm, ZRC 2004.0546, 24825.009N
124809.869E, fast flowing tributary of Nagura River, below
a reservoir, Ishigaki Island, Ryukyu Islands, pH 7.2, coll. Y.
Cai and T. Naruse, 13 Jun 2000; 22 males, cl 3.0-4.1 mm,
7 females, cl 3.2-4.3 mm, ZRC 2004.0547, 24816.609N
123852.749E, river mouth at Haemida beach, Iriomote
Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng and T.
Naruse, 14 Jun 2000; 6 males, cl 4.0-4.5 mm, 2 females, cl
3.5-4.0 mm, 6 ovigerous females, cl 5.4-6.4 mm, ZRC
2004.0548, 4816.649N 123852.809E 2, shallow freshwater
stream with pH 6.8, Aira River, Iriomote Island, Ryukyu
Islands, coll. Y. Cai, N. K. Ng and T. Naruse, 14 Jun 2000;
7 males, cl 3.1-4.7 mm, 1 ovigerous female, cl 6.2 mm, ZRC
2004.0549, 24816.609N 123852.749E, shallow freshwater
stream, downstream of Aira River, Iriomote Island, Ryukyu
Islands, coll. Y. Cai, N. K. Ng and T. Naruse, 14 Jun 2000;
1 male, cl 5.2 mm, 2 females, cl 4.9-5.2 mm, ZRC,
24818.249N 123851.219E, very small, shallow stream, up in
the hills, headwater of Nakama River, Iriomote Island,
Ryukyu Islands, pH 7.4, coll. Y. Cai, N. K. Ng and T.
Naruse, 15 Jun 2000; 6 males, cl 4.8-5.3 mm, 8 females,
cl 4.8-7.5 mm, 1 ovigerous female, cl 7.0 mm, ZRC
2004.0550, 24818.329N 123851.199E, freshwater, slow
flowing stream, slightly downstream of headwater of
Nakama River, Iriomote Island, Ryukyu Islands, pH 7.4,
coll. Y. Cai, N. K. Ng and T. Naruse, 15 Jun 2000; 35
males, cl 3.2-5.3 mm, 4 females, cl 5.4-6.3 mm, 15
ovigerous females, cl 5.5-7.2 mm, ZRC 2004.0551,
24818.399N 123851.299E, downstream from the headwater
of Nakama River, Iriomote Island, Ryukyu Islands, coll. Y.
Cai, N. K. Ng and T. Naruse, 15 Jun 2000; 26 males, cl 3.84.5 mm, 20 females, cl 3.3-5.6 mm, 3 ovigerous females, cl
5.4-6.1 mm, ZRC 2004.0552, 24823.659N 123851.849E, fast
flowing water, about 200 meters from sea, Omija River,
Iriomote Island, Ryukyu Islands, coll. Y. Cai, N. K. Ng and
T. Naruse, 15 Jun 2000; 8 males, cl 3.8-5.5 mm, 13 females,
cl 4.5-7.1 mm, 3 ovigerous females, cl 5.5-6.5 mm, ZRC
2004.0553, fast flowing water, upper part of Omija River,
Iriomote Island, Ryukyu Islands, coll. Y. Cai and T. Naruse,
15 Jun 2000; 9 males, cl 3.4-4.3 mm, 6 females, cl 4.1-4.6
mm, ZRC 2004.0554, 24823.659N 123851.849E, fast flowing water, about 200 metres from sea, Omija River, Iriomote
Island, Ryukyu Islands, coll. Y. Cai and T. Naruse, 16 Jun
2000; 15 males, cl 4.3-5.1 mm, 5 females, cl 3.3-7.3 mm, 8
ovigerous females, cl 6.3-7.1 mm, ZRC 2004.0555,
123851.299E 24818.399N, upstream of Nakama River, near
the head water, Iriomote Island, Ryukyu Islands, coll. Y. Cai
and T. Naruse, 16 Jun 2000; 2 males, cl 3.9-5.1 mm, 2
ovigerous females, 7.2-7.6 mm, egg 3.0 3 5.2 mm, ZRC
2004.0556, 24818.399N 123851.299E, headwaters of Nakama River, Iriomote Island, Ryukyu Islands, coll. Y. Cai
and T. Naruse, 16 Jun 2000; 43 males, cl 4.1-5.7 mm, 18
females, cl 3.8-7.5 mm, 1 ovigerous female, cl 7.4 mm,
CAI ET AL.: STIMPSON’S JAPANESE ATYIDS
ZRC 2004.0557, 24835.139N 124818.919E, pH 7.7, a stream
in Hirakubo, Ishigaki Island, Ryukyu Islands, coll. Y. Cai,
N. K. Ng and T. Naruse, 17 Jun 2000; 1 male, cl 2.9 mm, 1
female, cl 5.1 mm, 3 ovigerous females, cl 5.3-6.8 mm,
ZRC 2004.0558, 24817.819N 124833.769E, pH 7.7, Kara
Stream, Ishigaki Island, Ryukyu Islands, coll. Y. Cai, N. K.
Ng and T. Naruse, 17 Jun 2000; 20 males, cl 3.6-5.7 mm, 7
females, cl 5.6-7.5 mm, 4 ovigerous females, cl 5.4-6.1 mm,
ZRC 2004.0559, 24824.369N 124814.829E, small stream in
Ohno town, Ishigaki Island, Ryukyu Islands, coll. Y. Cai, N.
K. Ng and T. Naruse, 17 Jun 2000; 4 males, cl 3.4-4.2 mm,
2 females, cl 5.0-7.8 mm, 3 ovigerous females, cl 6.7-7.5
mm, ZRC 2004.0560, Okuma River, Okinawa Island,
Ryukyu Islands, coll. P. K. L. Ng et al., 31 Jun 1998; 1
male, cl 4.8 mm, 1 ovigerous female, cl 8.0 mm, ZRC
2004.0561, Taiho River, Okinawa Island, Ryukyu Islands,
31 May 1998; 5 males, 4.4-5.2 mm, 5 females, cl 6.9-7.9
mm, 4 ovigerous females, cl 6.8-7.2 mm, NSMT-Cr 1226,
Akina-gawa River, Amami-oshima Island, 17 July 1988,
coll. M. Takeda; 3 females, cl 3.0-3.2 mm, 4 males, cl 4.44.7 mm, NSMT, upstream of Nakama river, Iriomote,
Ryukyu Islands, M. Tomokuni, 3 Nov 1985; 2 males, cl 5.35.6 mm, 4 females, cl 6.6-7.3 mm, NSMT, Kiyose-bashi
bridge, Chichi-jima Island, Ogasawara Islands, 13 Oct 1974;
1 female, cl 7.7 mm, NSMT, upper area of Kawauchi-gawa
River, Amami-oshima Island, Ryukyu Islands, coll. M.
Takeda, 15 Jul 1988; 2 males, cl 5.0-5.2 mm, UR1176,
Sueyoshi, Hachijo-jima Island, Izu Islands, 28 Jun 1976;
2 males, cl 5.0-5.4 mm, 1 female, cl 6.7 mm, 1 ovigerous
female, cl 6.8 mm, UR1114, Kawauchi-gawa River,
Amami-oshima Island, Ryukyu Islands, coll. M. Takeda,
18 Jul 1988; 3 males, cl 4.0-4.4 mm, 2 females, cl 4.6-4.8
mm, 4 juv., UR1904, from an outlet stream of the
Isunadamari Pond, Kikai-jima Island, coll. Nagaoka, 26
Oct 1993; 6 females, cl 4.5-6.7 mm, 4 males, cl 4.8-5.5 mm,
UR1903, Kikai-jima, Bird Hill Park, coll. Nagaoka, 27 Oct
1993; 4 males, cl 5.5-5.8 mm, 3 ovigerous females, cl 6.58.7 mm, UR1905, Kikai-jima Island, pond at edge of
stream, coll. Nagaoka, 26 Oct 1993; 1 male, cl 6.0 mm,
UR1761, a substream of Nakama River, Iriomote Island,
Ryukyu Islands, Aug 1968; 1 female, cl 5.4 mm, UR1230,
Sumiyoh-gawa River, Amami-ohshima Island, Ryukyu
Islands, coll. M. Takeda, 17 Jul 1988; 1 male, cl 4.8 mm,
1 ovigerous female, cl 6.9 mm, UR1112, Shino-kawa River,
Amami-oshima Island, Ryukyu Islands, coll. M. Takeda, 17
Jul 1988; 4 females, cl 5.4-6.8 mm, UR1182, Kauauchigawa River, Amami-ohshima Island, Ryukyu Islands, coll.
M. Takeda, 15 Jul 1988; 3 males, 4.9-5.9 mm, 4 females, cl
6.5-7.5 mm, 2 ovigerous females, cl 6.1-6.5 mm, UR1218,
Kawauchi-gawa River, Amami-ohshima Island, Ryukyu
Islands, coll M. Takeda, 18 Jul 1988; 1 male, cl 4.2 mm, 2
females, cl 4.8-5.2 mm, 2 ovigerous females, cl 5.2-5.6 mm,
UR1135, Kanyu-gawa River, Amami-ohshima Island,
Ryukyu Islands, coll. M. Takeda, 16 Jul 1988; 2 ovigerous
females, cl 4.3-5.2 mm, UR1759, no data.
Comparative Material Examined.—34 males, cl 3.5-4.6 mm,
9 females, cl 3.2-5.0 mm, 4 ovigerous femals, cl 4.3-6.3
mm, 39 juveniles, syntypes of Caridina typus brevirostris
417
Roux, 1934, SMF-2148, Elat and Ohoinaugau, Kei Islands,
Indonesia, coll. Merton, 3 Jun 1908.
Description.—Rostrum reaching near to end of basal
segment, or to end of second segment of antennular
peduncle, unarmed dorsally, armed ventrally with 1-4 teeth;
antennal spine fused inferior orbital angle; pterygostomian
margin subrectangular. Sixth pleomere 0.40 times as long
as carapace, 1.2 times as long as fifth somite, shorter than
telson. Telson 2.8 times as long as wide, terminating in
posteromedian projection; 4 or 5 pairs of dorsal spinules,
situated on distal two-third of telson length, 1 pair of
dorsolateral spines near distal end, 4 or 5 pairs of spines on
distal margin, lateral pair subequal to intermedian pairs;
preanal carina lacking spine.
Eyes well developed. Antennular peduncle slender, 0.7
times as long as carapace; basal segment half length of the
antennular peduncle, second segment 1.5 times as long as
third one; stylocerite reaching 0.8 times length of basal
segment. Scaphocerite 3.3 times as long as wide.
Mouthparts similar to C. grandirostris. Third maxilliped
reaching to end of second segment of antennular peduncle,
with a short exopod, reaching to middle of antepenutimate
segment, ultimate segment distinctly shorter than penultimate segment.
Epipods on first 4 pereiopods. First pereiopod short,
reaching to end of basal segment of antennular peduncle,
merus 2.2-2.8 as long as broad, slightly longer than carpus;
carpus excavated strongly anteriorly, shorter than chela, 1.21.6 times as long as high; chela 2.0-2.2 times as long as
broad; length of fingers variable, at most shorter than palm,
sometimes as long as or even slightly longer. Second
pereiopod reaching end of second segment of antennular
peduncle, merus shorter than carpus, 4.3-5.5 times as long
as broad; carpus distinctly longer than chela, 5.0-6.2 times
as long as high; chela 2.7-3.0 times as long as broad; fingers
1.3-1.8 times as long as palm. Third pereiopod long,
slender, reaching to end of antennular peduncle, propodus
distinctly shorter than merus, 7.7-9.1 times as long as broad,
3.7-4.4 times as long as dactylus; dactylus ending in a strong
claw; 2.4-2.9 times as long as wide (spines included), flexor
margin with 4-6 accessory spines strongly curved inwards;
fifth pereiopod reaching to end of scaphocerite, propodus
distinctly longer than merus, 11-14 times as long as broad,
3.3-3.7 times as long as dactylus; dactylus 3.3-4.2 times as
long as wide, with 60-77 spinules of flexor margin.
Endopod of male first pleopod extending to half length of
exopod, elongate, subrectangular, 2.0-2.3 times as long as
broad, with distinct appendix interna near distal end of
endopod. Appendix masculina of male second pleopod
slender, reaching to 2/3 length of endopod, inner and distal
surface densely lined with long spines; appendix interna at
basal 1/3 of appendix masculina, extending to distal 1/3 of
appendix masculina. Uropodal diaeresis with 19-24 spinules.
Egg size 0.45-0.48 3 0.26-0.23 mm in diameter.
Remarks.—Bouvier (1925) treated C. exilirostris Stimpson,
1860, as a junior synonym of C. typus. Chace (1997) recently
commented that Caridina exilirostris is possibly a synonym
of C. valladolidi Blanco, 1939. Cai and Ng (2001) com-
418
JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 26, NO. 3, 2006
pared Caridina valladolidi Blanco, 1939, with C. exilirostris, and stated that Caridina exilirostris is most probably
C. typus. The rostrum of Caridina exilirostris as described
by Stimpson (1860:98) was as follows ‘‘. . . valde gracile,
compressum, angustum, acutum, medium articuli penultimi
antennularum pedunculi parce superans . . .’’ (slender,
compressed, narrow, acute, reaching to middle of second
segment of antennular peduncle), while that of C.
villadolidi, as a rule, reaches to or nearly to the end of the
antennular peduncle. Until now, C. villadolidi has never
been reported from the Ryukyu Islands, the type locality of
C. exilirostris. Stimpson (1860) noted the similarity between
C. typus and C. exilirostris, but failed to recognize that both
are identical. To stabilize the taxonomy of these species, an
ovigerous female, cl 7.4 mm, Okuma River, Okinawa Island
(type locality) is hereby designated as the neotype (NSMT)
of C. exilirostris Stimpson, 1860.
That the length of the rostrum of C. typus is variable is
well known (see Ng, 1995). During a faunistic survey by K.
Satake on the freshwater shrimps of Chichi-jima Island and
Haha-jima Island, Ogasawara Islands, 67 lots of specimens
were collected, but only three of them have a very short
rostrum which does not reach beyond the end of the
antennular peduncle. All these specimens were found from
the headwaters. The specimens found at the headwater
region of the Nakama River also have relatively shorter
rostrums when compared with those living in the lower parts
of the same river. Bouvier (1925: 126, fig. 271) reported
C. typus var. acuminata from Poulo Condor (So’n Island),
southern Vietnam. These specimens (ca. 30 specimens in
bad condition, MNHN Na-932) were re-examined and
shown to be C. typus, which have a relatively shorter
rostrum. The syntypes of Caridina typus brevirostris Roux,
1934, were reexamined. Caridina typus brevirostris has
a short rostrum, very short fingers of the first pereiopod, and
sexually dimorphic dactylus of third and fourth pereiopods.
It should be regarded as a distinct species. As the name
Caridina typus brevirostris Roux, 1934, is preoccupied by
C. brevirostris Stimpson, 1860, a replacement name has to
be proposed. This matter is now being investigated by the
first author.
Habitat.—Caridina typus occurs in rivers or streams on
islands or in coastal areas, prefers to hide under rocks or
stones during the day time and go out to forage at night.
ACKNOWLEDGEMENTS
YC would like to thank Prof. Masatsune Takeda (NSMT), Drs. Charles
Fransen (RMNH), Nguyen, Ngoc-Ho (MNHN), Michael Türkay (SMF),
Ambro Hänggi (NHMB), Dirk Platvoet (AMZ) and Tohru Naruse (UR) for
their hospitality when he visited their museums and laboratories and for the
loans of specimens for his study; for Dr. Junji Okuno (Coastal Branch of the
Natural History Museum and Institute, Chiba, Japan); Dr. Hiroshi Suzuki
(Faculty of Fisheries, Kagoshima University, Shimoarata, Kagoshima,
Japan) for sending their collections for this study. KS would like to thank
Mr. R. Ueno (NIES) for his encouragement and collaboration in the field,
Dr. Machiko Nishino (Lake Biwa Museum), Dr. Seiichi Nohara, Dr.
Toshiki Natori, Dr. Tohru Yabe (NIES), and Dr. Hirokatsu Utagawa (Japan
Society for the Promotion of Science) for their discussions, Dr. Kazuo
Horikoshi, Mr. Makoto Inaba and Mr. Hajime Suzuki (Institute for
Boninology) for their help and guidance in Chichi-jima, Mr. Hayato Chiba
(Ogasawara Village Office) and Mr. Yoshio Hoshi (Villa Kobunoki) for
their help and guidance in Haha-jima.
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RECEIVED: 12 November 2004.
ACCEPTED: 20 January 2006.
