Source : MNHN, Paris

Source : MNHN, Paris

Source : MNHN, Paris

Source : MNHN, Paris

Source : MNHN, Paris

Source : MNHN, Paris

Il

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’J MÉMOIRES _

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MUSÉUM NATION AL

D'HISTOIRE NATURELLE

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Source : MNHN, Paris

Source : MNHN, Paris

MEMOIRES

DU

MUSÉUM NATIONAL

D'HISTOIRE NATURELLE

NOUVELLE SÉRIE
Série A, Zoologie
TOME XXXIII

PARIS

ÉDITIONS DU MUSÉUM
38, rue Geoffroy-Saint-Hilaire (V e )

1965

Source : MNHN, Paris

Source : MNHN, Paris

TABLE DES MATIÈRES

Fascicule 1

(1965)

Pages

L. B. Holthuis, The Atyidae of Madagascar. 1-18

Fascicule 2

(1965)

E. Fischer-Piette et J. Bédoucha, Mollusques terrestres

operculés de Madagascar. 49-92

Fascicule 3

(1965)

Ch.-P. Blanc, Études sur les Iguanidae de Madagascar . . . 93-148

Fascicule 4

(1965)

M. Gabe et H. Saint-Girons, Contributions à la morphologie
comparée du cloaque et des glandes épidermoïdes de la

région cloacale chez les lépidosauriens. 149-292

Source : MNHN, Paris

Source : MNHN, Paris

n v

MÉMOIRES

DU

MUSÉUM NATIONAL

D’HISTOIRE NATURELLE

NOUVELLE SÉRIE

Série A, Zoologie

TOME XXXIII
FASCICULE I

L. B. HOLTHUIS

X THE ATYIDAE OF MADAGASCAR

PARIS

ÉDITIONS DU MUSÉUM
38, rue Geofïroy-Saint-Hilaire (V e )

1965

Source : MNHN, Paris

*

Source : MNHN, Paris

J T-

b U

MÉMOIRES Mi MISITtl NATIONIl U'HISTOIRE NATURELLE

Série A, Zoologie. Tome XXXIII, Fascicule 1. — 1965

, THE ATYIDAE OF MADAGASCAR

L. B. HOLTHU1S

Rijksmuseum van Natuurlijke Historié, Leiden

The présent paper is based in the first place on an extensive collection
ol Atyidae from ail over Madagascar which is the property of the Institut
de Recherche Scientifique de Madagascar at Tananarive ; this material
was placed at my disposai by Dr. R. Paulian, formerly deputy director of
that Institute, for which I wish to tender liim my deeply felt gratitude.
Also incorporated in this report is the important collection of Madagascar
Atyidae brought together during the ‘Oesterreichischen Madagaskar-
Expedition 1958”, which was entrusted to me for study by the leader of the
expédition, Dr. F. Starmühlner of the Zoological Institute of Vienna
University to whom I like to express my sincere thanks. The few specimens
of Madagascar Atyidae présent in the Leiden Muséum are also taken into
account. Just before the manuscript was finished a small but interesting
collection of Madagascar Atyidae was received by the Leiden Muséum.
For this collection, which is also dealth witli here, I am most indebted to
Mr. Y. Therezien, ingénieur-hydrobiologiste of the Centre Technique Fores¬
tier Tropical of Tananarive. The Madagascar Institute donated duplicates
of its collection to the Leiden Muséum, which obtained also the entire
collection of the Austrian Expédition.

In order to make this paper of more use to those workers in Madagascar
interested in the Atyid fauna of the island, ail species so far reported from
Madagascar and not represented in the above collections hâve been listed
here also. In this way the présent report gives a complété review of the
actual State of our knowledge of this group. In the synonymy of each species
a reference to the original description is given, as well as references to ail
Madagascar records of the species that are known to me ; also a full énumé¬
ration of the localities from where the species has been reported is provided.

At présent 5 généra and 23 species of Atyidae are known from Madagascar.
The généra may be distinguished with the help of the following key :

Mémoires du Muséum. — Zoologie, t. XXXIII. 1

Source : MNHN, Paris

p. . _ MaD of Madagascar, showing the more important localitics mentioned ln the
t»vt Insert showing the situation of lîuropa Island and the division of Madagas-
car in to provinces (I, Diégo-So.rez ; II, Mojanga , 111 Taj.narlvc ; IV, Tnléar ; V.
Tamatave ; VI, Fianarantsoa). Scale of large map 1 : 7.500.000.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

1. Exopods on ail pereiopods. Eyes degenerate, cornea absent or strongly

— reduced. Rostrum short, slender, without teeth. 2

Exopods absent from ail pereiopods. Cornea well developed or reduced,
always with pigment. Rostrum with or without teeth . 3

2. Eyes without any pigment, bulletshaped. Pterygostomian spine absent.

Second maxilliped without podobranch. Endopod of first male pleopod
with appendix interna . Typhlopatsa

— Eyes with a strongly reduced poorly pigmented cornea placed on the

antérolatéral angle of the broad peduncle. Eye broad, quadrangular,
excavate anteriorly. Second maxilliped with podobranch. Endopod
of first male pleopod tapering to a narrow elongate tip, without appendix
interna. Antecaridina

3. First pereiopod without arthrobranch. Cornea reduced or well developed
. Parisia

— First pereiopod with arthrobranch and pleurobranch. Cornea always

well developed. 4

4. Carpus of second leg elongate, more than three times as long as high,

not deeply excavated. Chelae always with a well developed palmar
portion . Caridina

— Carpus of second leg less than twice as long as high, deeply excavated.

Dactylus of cheliped usually as long as propodus and articulating at
base of propodus so that there is no palmar portion. Alya

Genus TYPHLOPATSA Holthuis, 1956

Only one species known :

Typhlopatsa pauliani Holthuis

Typhlopatsa pauliani Holthuis, 1956, p. 53.

Typhlopatsa Pauliani Holthuis, 1956a, p. 98, fig. 1.

For the description of the species I may refer to the second of the above
cited references.

The species is cavernicolous and so far was found only in the Mitoho Cave,
NE corner of Tsimanampetsotsa Lake, Mahafaly Province, SW Madagascar.

Genus ANTECARIDINA Edmondson, 1954

The genus is monotypic, the only species is :

Source : MNHN, Paris

L. B. HOLTHUIS

Antecaridina lauensis (Edmondson) (fig. 2)

Mesocaris lauensis Edmondson, 1935, p. 13, fig. 4.

Madagascar Institut!-: :

— Europa Island, W of Madagascar ; saline pool ; June 1951 ; R. Paulian. — 12 spe-

The specimens are 10 to 15 mm long.

The rostrum is unarmed above and below, it is rather slender and reaclies
about to the middle of the second segment of the antennular peduncle.
In dorsal view it is elongately triangular. There is no supra-orbital spine.
The antennal spine is placed on the lower orbital angle, it is well developed
and sharp. A conspicuous pterygostomian tooth is présent.

The telson dorsally bears none to two, generally two pairs of spinules.
The posterior margin of the telson is rounded, bears a short outer pair
of spinules, a latéral pair which is more than 5 times as long as the outer
pair and 3 to 5 intermediate spines, the outer of which are longest.

The eyes hâve the comea strongly reduced, forming only a small rounded
poorly pigmented swelling on the outer antérolatéral corner of the peduncle,
which itself is distinctly shorter than broad.

The segments of the antennular peduncle are short. The stylocerite
is long and slender and reaches distinctly beyond the base of the second
segment. The antérolatéral angle of the first segment is only slightly produced.

The scaphocerite is oval, with a short final tooth on the outer margin.
This tooth is far outreached by the lamella. At the base of the scaphocerite
the antennal peduncle shows a strong spine.

The mouth parts are of the normal type. They are flgured here.

The branchial formula runs as follows :

MAX II. LIFE DS

1 2 3

Pleurobranchs
Arthrobranchs
Podobranchs
Epipods
Exopods

PER El O PO DS

2 3 4

The first pereiopod lias the fingers about as long as the palm. The carpus
is deeply hollowed anteriorly and is more than twice as long as high, shorter
than the chela and somewhat longer than the merus. The second pereiopod
is more slender and longer than the first. The fingers are longer than the
palm. The carpus is fully 4 times as long as high and much longer than eithcr
chela or merus. In the third leg the dactylus ends in two claws, while three
slender spines are placed on the posterior margin. The propodus is more
than 2.5 times as long as the dactylus. The carpus bears a distal movable
spine, the merus lias a similarly situated spine and 4 movable spines on the
posterior margin ; a movable spine may also be seen on the ischium. In the
fifth leg the dactylus also ends in two claws, on the posterior margin there

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

5

. 2. — Anlecaridina lauensis (Edmondson). a, anterior part of body in latéral view ;
b, anterior part of body in dorsal view ; c, tip of telson ; d, eye ; e, /, mandible ;
g, maxillula ; h, maxilla ; i, flrst maxilliped ; /, second maxilliped ; k, third maxilliped ;
l, flrst pereiopod ; m, second pereiopod ; n, third pereiopod ; o, fifth pereiopod ;
p, endopod of flrst pleopod of male ; q, appendix interna and appendix masculina
of second pleopod of male ; r, outer part of diaeresis of uropodal exopod. a, b, k-o, x 16;
c-j, p-r, x 33. Ali ligures after specimens from Europa Island.

Source : MNHN, Paris

L. B. HOLTHUIS

is a row of short hairs. The propodus is slightly more than twice as long as
the dactylus. Both merus and carpus hâve a distal movable spine, no other
such spines are seen here.

The endopod of the first pleopod of the male is oval, tapering to a slender
elongate tip. The appendix masculina of the second male pleopod is much
shorter than the very slender appendix interna, il bears a tuft of strong
setae on the top. The outer margin of the exopod of the uropods ends in a
small tooth, to the inside of which a slender movable spine is placed ; there
are no other spines on the diaeresis.

Colour. Edmondson described the species as being bright red when alive.
No information about the colour of the Madagascar specimens is available.

The species so far was known only from two localities in the Lau Islands,
Fiji Group, Polynesia, viz., Numbu Cave on Namuka Island, and Wangavu
Island. The water in Numbu Cave is brackish, on Wangavu Island the species
was found in a salty lake. The présent record of the species from a sait
water pool on Europa Island confirms the preference for sait water of this
species, an unusual feature in the présent family which always is considered
to be a typical freshwater group.

The tact that the species is found in localities geographically so widely
apart as Europa Island and the Fiji Archipelago makes it very probable
that it will be found also in the intermediate area (1).

Rathbun (1906, p. 919, fig. 67) dealt with small vermillion Atyids
found "in small fresh or slightly brackish water pools in lava flow, near
sea” 5 miles south of Puako Bay near Hilo on the island of Hawaii. Judging
by her account the specimens are extremely close to Antecaridina lauensis,
dilfering mainly by the shorter rostrum and the less slender second pereiopod.
The absence of a stylocerite in Rathbun’s drawing evidently is an omission
by the artist. Without reexamination of Rathbun’s material it is impossible
to place the species, but it does not seem probable that her identification
with Caridina bievircstris Stimpson is correct, the latter is a tiue freshwater
form, probably belonging to the genus Caridina (1).

Antecaridina Edmondson forms with the généra Typhlalya Creaser,
Spekocaris Matjasiô, Typhlopatsa Holthuis and Slygiocaris Holthuis, a
homogeneous section of the family Atyidae, quite distinct from Bouvier’s
(1925, pp. 40-42) série paratyienne, série caridellienne and série caridinienne ;
it could well be named série typhlatyienne.

(1) Note with the correction : since the abovc was written, a paper, entitled « on red
coloured shrimps (Deeapoda, Caridea) from tropical land-locked saltwatcr pools », was
published (Holthuis, 1963, 7ool. Meded. Leiden, vol. 38, pp. 261-279, figs. 1, 2) ; in this
publication Antecaridina lauensis is reported from the Southern Red Sea, while some
additional information on the habitat of the présent Madagascar specimens of thaï species
is providcd. In the sanie paper Rathbun’s » Caridina breuirostris . is described as a new
genus and new species under the name Halocaridina rubra.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

Genus PARISIA Holthuis, 1956

The genus consists of three species, ail of which inhabit Madagascar.

They may be distinguished as foUows :

1. Eyes strongly reduced with only a small pigment spot. Antennal spine

merged with the lower orbital angle. 2

— Eyes with the cornea well developed and rounded, provided with dark

pigment. Antennal spine distinctly separated from the lower orbital
angle. Rostrum well developed, with teeth on both upper and lower
margins . macrophthalma

2. Rostrum well developed, reaching about to the end of the antennular
peduncle, with teeth on both upper and lower margins .... microphlhalma

— Rostrum short, not reaching beyond the eyes, without either dorsal

or ventral teeth . edentata

Parisia edentata Holthuis

Parisia edentata Holthuis, 1956, p. 55 ; Holthuis, 1956a, p. 105, fig. 3.

For the description of the species I refer to the last mentioned reference.
The species is only known from the Southern part of the Antsingy Mts.
between the Manambolo and Morafenobé rivers near Bekopaka, Majunga
province, W Madagascar ; in a very dark deep well.

Parisia microphthalma (Fage)

Caridina microphthalma Fage, 1946, p. 324, figs. 1, 2.

Parisia microphthalma Holthuis, 1956, p. 54 ; Holthuis, 1956a, p. 103, fig. 2.

Descriptions and figures of the species hâve been given by Fage (1946)
and Holthuis (1956a), to which I may refer here.

So far the species has been collected only in the Grotte des Fanihy,
Ankarana Mts., N of Ambilobé, N Madagascar, in a fresh water pool in
total darkness.

Parisia macrophthalma Holthuis

Parisia macrophthalma Holthuis, 1956, p. 55 ; Holthuis, 1956a, p. 107, fig. 4.

For a description and figures of the species I refer to the last cited
reference.

Source : MNHN, Paris

L. B. HOLTHUIS

The species was found together with the previous one in the Grotte des
Fanihy, Ankarana Mts., N of Ambilobé, N Madagascar, in a fresh water
pool in total darkness.

Genus CARIDINA H. Milne Edwards, 1837

The genus Caridina, which in Madagascar is represented by at least
16 species, is one of the notoriously diiïicult généra of Decapoda Macrura.
Even Bouvier’s (1925) monograph of the Atyidae did not put an end to
these difficulties. Several of the characters used by Bouvier to distinguish
between his species (and subspecies) prove to be too variable to be of great
value ; this is especially true of most of the relative measurements. To
several other characters too little importance was attached by Bouvier.

As is clearly shown by the présent material, local populations may show
a remarkable constancy in certain characters, while if représentatives of
various populations of a whole species are considered, these characteis
vary greatly, often linking two forms, which at first sight seem to represent
distinct species. For this reason an exact knowledge of the various species
of Caridina of Madagascar can only be obtained by a zoologist who is sta-
tioned in Madagascar and can explore the fauna of ail the different river
Systems and study the influence of locality, environment and âge on the
various characters. Only in this way a coirect idea of the species, their
variability and their mutual relations can be obtained.

It is the purpose of the piesent paper, to place in the hands of such a
zoologist ail the data concerning our présent knowledge of the Madagascar
Atyidae so that it may serve as a foundation on which a final révision of
this fauna can be based.

It will be of the utmost importance to known the changes which each
species undergoes in growing up. The material at hand shows that there
are considérable differentes between juvénile and adult specimens, but
the actual relation is not sufficiently well known. Also the influence of the
habitat on certain characters is important. So Gordon (1930) pointed out
that in lacustrine specimens of C. nilotica the rostrum and also the appcn-
dages are more slender than in specimens from shore collections. As in most
instances little or nothing is known about the conditions under which the
présent material lived, hardly anything could be done in this respect. Both
the inter- and the intra-populational variation should be given ail attention
in order to make a solution of the species problem of the Madagascar Cari¬
dina possible. I fully realize that my présent attempts in this direction
are very feeble, and though I am sure that several of my conclusions will
prove to be incorrect, 1 présent these my results with the hope that they
will be a stimulus to someone with more favourable facilities to untie the
numerous knots that still are left (or perhaps even added to) the systematics
of the Madagascar Caridina species.

The following key to the Madagascar species of Caridina is only intended
as a first attempt. Especially the last part (7 and beyond) should be used
with some reserve as the variability of the various characters employed

Source : MNHN, Paris

THE ATY1DAE OF MADAGASCAR 9

is rather great. C. brevirostris brevipes is not inserted in this key as too
little is known of this form.

1. Endopod of first pleopod of male with appendix interna.2

— Endopod of first pleopod of male without appendix interna ... 6

2. Upper border of rostrum without teeth. Rostrum short, not reaching
beyond second segment of antennular peduncle. Intermediate spines
on posterior margin of telson longer tlian the latéral pair. Antennal
spine placed on the lower orbital angle. typus

— Upper border of rostrum with teeth. Intermediate spines on posterior
margin of telson as long as or shorter than the latéral pair ... 3

3 .Rostrum straight and short, not reaching beyond the second segment
of antennular peduncle. Distal part of upper margin of rostrum unarmed,
no subapical teeth présent. Antennal spine placed on lower orbital
angle. japonica

— Rostrum with the distal part curved up, reaching distinctly beyond

second segment of antennular peduncle. Distal part of upper margin
of rostrum unarmed apart from one or more subapical teeth. Antennal
spine placed below the very distinct lower orbital angle.4

4. Posterior margin of telson ending in a médian point. Preanal carina

with a sharp posteriorly directed spine. Eggs numerous and small
(0.2 — 4.0 X 0.3 — 0.6 mm).5

— Posterior margin of telson rounded. Preanal carina blunt without a
spine. Eggs fewer and larger (0.5 — 0.6 x 0.7 — 1.0 mm) .. nilotica

5. Dorsal margin of rostrum with 13 to 23 spines in basal part, which

are placed close together . longirostris

— Dorsal margin of rostrum with 5 to 13 spines in the basal part, which

are placed widely apart. graciliroslris

6. 6 to 8 spines of the dorsal margin of the rostrum placed behind the orbit.

Stylocerite reaching distinctly beyond the base of the second segment
of the antennular peduncle. Intermediate spines of posterior margin
of the telson larger than the latéral pair. serratirostris

— 0 to 6 spines of the dorsal margin of the rostrum placed behind the
orbit. Stylocerite at most attaining the end of the first segment of the

antennular peduncle.7

7. Intermediate spines of the posterior margin of the telson longer than

the latéral pair, with a chitine plug in the middle.8

— Intermediate spines of the posterior margin of the telson shorter than
the latéral pair.10

8. Anterior margin of carpus of first cheliped deeply excavated ; this

carpus less than twice as long as wide, shorter than the chela. Lower
margin of rostrum with about 8 teeth ; upper margin with 0 to 16

. angulata

— Anterior margin of carpus of first cheliped not deeply excavated, carpus

itself more than twice as long as wide, longer than the chela ... 9

9. Upper margin of rostrum with 10 to 13 teeth. petiti

— Upper margin of rostrum without teeth, seldom with 1, rarely with 7
. calmani

Source : MNHN, Paris

10

L. B. HOLTHUÏS

10. Dactylus of third pereiopod with 15 to 17 teeth on posterior margin...
. edulis

— Dactylus of third pereiopod with 3 to 7 teeth on posterior margin. 11

11. Upper margin of rostrum without teeth, rostrum short.12

— Upper margin of rostrum with teeth.13

12. Diaeresis with 7 to 13 spinules. isaloensis isaloensis

— Diaeresis with 20 to 23 spinules. lamiana

13. 4 to 6 of the dorsal teeth of the rostrum placed behind the orbit. Poste¬
rior margin of telson stepped. troglophila

— 0 to 3 of the dorsal teeth of the rostrum placed behind the orbit . 14

14. Rostrum in adult specimens very long, reaching much beyond the

antennular peduncle. Ultimate part of upper margin of rostrum without
teeth. xiphias

— Rostrum in the adults shorter than the antennular peduncle or, if longer,

with a subapical tooth.15

15. Rostrum in adults usually not reaching beyond the base of the third

antennular segment, evenly dentate or with a short distal unarmed

space.16

— Rostrum as long as or longer than antennular peduncle, with a distinct

unarmed stretch behind the subapical tooth. norvestica

16. Dorsal teeth of rostrum very high and strong. hova

— Dorsal teeth of rostrum small. isaloensis grandidieri

Caridina typus H. Milne Edwards (fig. 3)

Caridina typus H. Milne Edwards, 1837, p. 363, pl. 25bis, figs.’4,5 ; Coutière,
1900, p. 1267 ; Bouvier, 1904, p. 134.

Caridina typa Bouvier, 1905, p. 88.

Caridina typus typica Bouvier, 1925, p. 250, figs. 272-295.

Madagascar Institute :

— Ankavandra on Manambolo River, W Madagascar ; July 1949 ; R. Paulian_

1 specimen.

— Betsingily on Manambolo River, W Madagascar ; July 1949 ; R. Paulian. —
7 specimens.

— Mahilaka, Bay of Ampasindava opposite Nosy Bé, NVV Madagascar ; 1946;
J. Millot. — 3 specimens.

— Bas-Sambirano near Ambanja, mainland opposite Nosy Bé, NW Madagascar;
swamp ; June 1948 ; J. Millot. — 4 specimens.

— Orangea near Diégo-Suarez, N Madagascar ; cave ; 1946; J. Millot. — 5 specimens
(2 ovigerous).

— Tamatave, F.. Madagascar ; fresh water ; April 1951; R. Paulian. — 8 specimens.

Austrian Madagascar Expédition :

Djabalabe rivulet, Nosy Bé ; 18 and 20 August 1958 ; F.M. 157, 163. — 17 spe¬
cimens.

— Djabala rivulet, Nosy Bé ; 22 August 1958 ; F.M. 171. — 2 specimens.

— l’asandava rivulet, Nosy Bé ; 16 August 1958 ; F.M. 145. — 48 specimens.

— Andranobe rivulet, Nosy Bé ; 17 August 1958 ; F.M. 119. — 9 specimens.

— Anjiamarango rivulet, Nosy Bé ; 21 August 1958; F.M. 168. — 9 specimens.

_ Amporaha rivulet, Nosy Bé ; 16 August 1958 ; F.M. 146a. — 6 specimens.

Source : MNHN, Paris

E ATYIDAE OF MADAGASCAR

Muséum Leiden :

— Andrarony River, Antalaha district, Diégo-Suarez province, N E Madagascar ;
8 km from the mouth of the river ; 7 February 1962; native name « orangaratra * ; Y. The-
rezien no. 34. — 3 specimens (2 ovigerous).

— Ambodiriana, Tamatave district, Tamatave province, E Madagascar; July 1957;
native name « orana » ; Y. Therezien no. 2. — 4 females (3 ovigerius).

The largest specimen is 36 mm long ; the ovigerous females measure
24 to 36 mm.

The rostrum of this species is short, in the présent material it reaches
to the base or almost to the end of the second segment of the antennular
peduncle. The upper margin is entire, the lower margin bears 1 to 3 (rarely
up to 6) ventral teeth.

The antennal spine is placed on the lower orbital angle and is completely
fused with it. The pterygostomian angle is recta ngularly rounded.

Fio. 3. — Caridina typus H. Milne Edwards, from Andranobe rivulet, Nosy Bé. a, anterior
part of body in latéral view ; b, tip of tclson ; c, pre-anal carina ; </, third maxillipcd ;
e, first perciopod ; /, second pereiopod ; g, third pereiopod ; h, fifth pereiopod ;
i, endopod of first pleopod of male ; /, outer part of diaeresis. a, d-h, x 16 ; b, c.

Source : MNHN, Paris

12

L. B. HOLTHUIS

The dorsal surface of the telson bears 5 or 6 rarely 4 pairs of spines. The
posterior margin of the telson ends in a small sharp point, the postérolatéral
angles are somewhat produced. The outer spines of the posterior margin
are placed somewhat over the latéral spines and are very short. The latéral
spines are long (more than 5 times as long as the outer spines) and strong.
The intermediate spines are 4 to 9 in number and are slightly longer than
the latéral spines, they are divided in two by a kind of chitin plug.

The eyes are well developed.

The stylocerite reaches beyond the middle of the basal segment of the
antennular peduncle but fails to reach the end of the segment. The antéro¬
latéral angle of the segment is pointed and short and fails to reach the end
of the basal 1/4 of the second segment. There is a rather broad and sharp
tooth at the outer margin of the antennal peduncle near the base of the
scaphocerite.

Epipodites are présent on the first 4 pereiopods.

The carpus of the first chelipeds is about 1.5 times as long as high and
deeply hollowed anteriorly, it is about as long as the merus and shorter
than the chela. In the second leg the carpus is about 5 times as long as high,
not deeply excavated anteriorly and longer than either merus or chela ;
the Angers are much longer than the palm. The propodus of the third pereio¬
pods are about 4 times as long as the dactylus, the latter bears 5 or 6 teeth
on the posterior margin. The propodus of the 5th leg is 3 1/2 to 4 1/2 times
as long as the dactylus which is armed with numerous closely placed comb-
like arranged spinules.

The first pleopod of the male has a distinct appendix interna on the
endopod. In Bouvier’s (1925, p. 127, fig. 293) illustration of the endopod,
the appendix evidently is folded so that it obtains a rather peculiar shape
there ; as shown by the présent material the appendix is perfectly normal.
The appendix masculina of the second male pleopod is almost twice as long
and twice as broad as the appendix interna.

On the exopod of the uropods the diaeresis bears 20 to 23 spinules.

The eggs are numerous and small. In the présent specimens they measure
0.2 x 0.4 mm.

Caridina lypus, the type species of the genus, has a very wide distri¬
bution in the Indo-West Pacific area, ranging from East Africa to Japan
and Polynesia. The previous records from Madagascar are : Madagascar
(Coutière, 1900), Tuléar, SW coast (Bouvier, 1904, 1925), Diégo-Suarez,
NE coast (Bouvier, 1904, 1925), torrents of Antongil Bay, NE coast (Bou¬
vier, 1904, 1905, 1925), Fort-Dauphin, SE coast (Bouvier, 1904, 1925).

Caridina japonica De Man (fig. 4)

Caridina japonica De Man, 1892a, p. 261, pl. 9, fig. 7.

Muséum Leiden :

Ambodiriana, Tamatave district, Tamatave province, E Madagascar ; July 1957 ;
native mime < orana ■ ; Y. Therezien no. 2. 1 specimen.

Heforona River, Périnct région, Moramanga district, ramatave province ; 11 l'e-
bruary 1961 ; native namc . orana » ; Y. Therezien no. 10. — 1 specimen.

Source : MNHN, Paris

THE ATY1DAE OF MADAGASCAR

13

— Brook near thc city limits of Ifanadiana, Ifanadiana district, Fianarantsoa province,
E Madagascar ; 20 February 1962 ; native name . tsipitika • ; Y. Therezien no. 22. —
1 specimen.

— Tolongoina River, Fort-Carnot district, Fianarantsoa province ; 21 February 1962 ;
native name • patsa ■ ; Y. Therezien no. 18. — 1 specimen,

The specimens, ail of which are males, are 24 to 38 mm long.

The rostrum reaches somewhat beyond the base of the second segment
of the antennular peduncle. It is rather high and straight. The upper margin
bears 16 to 22 small movable teeth, ail of which are placed on the rostrum
proper ; they are close together and leave the ultimate 1/4 or 1/5 of the
upper margin unarmed ; no subapical teeth are présent. The first dorsal
tooth is placed over or slightly before the posterior limit of the orbit. The
lower margin of the rostrum bears 7 to 12 immovable, rather small teeth.
The latéral carina of the rostrum is quite distinct in the basal part and
merges with the orbital margin.

The antennal spine is placed on the lower orbital angle ; it is quite strong.
The pterygostomial angle is bioadly rectangularly rounded and liardly
forwards produced.

The sixth abdominal somite is about 0.5 to 0.6 times as long as the cara¬
pace. The telson bears 5 or 6 pairs of dorsal spinules, sometimes a seventh
spinule is présent on one side. The posterior margin of the telson ends in
a short médian tooth. The outer pair of posterior spinules is short, it does
not reach beyond the basal fourth of the latéral spines. There are 6 to 9
intermediate spines which are almost as long as the laterals ; they are
provided with fine setae. The preanal carina is produced to form a blunt-
topped triangular lobe ; it bears no spine.

The eyes are well developed with a globular well-pigmented cornea.
The antennular carina is dorsally produced to a high and bluntly topped
lobe. The antennula is short and robust with the various segments relatively
short and broad. The stylocerite is slender and sliarply pointed ; it reaches
beyond the middle of the basal antennular segment, but fails by far to
reach the end of it. The antero-lateral tooth of the basal segment is well
developed and measures about 1/4 of the length of the second segment. The
second segment is much shorter than the first and slightly longer than the
third. The outer margin of the scaphocerite is about straight and ends in a
sharp tooth which practically attains the end of the antennular peduncle ;
it is far overreached by the lamella. A sharp triangular tooth is présent
on the antennal peduncle near the external part of the base of the scapho¬
cerite.

The branchial formula is typical for the genus. Epipods are présent
on the first four pereiopods. The third maxilliped reaches to or with part
of the distal segment beyond the antennular peduncle. The length of the
ultimate segment is 4/5 of, or almost equal to that of the penultimate ;
it ends in a dark tooth and bears some spines in the distal part of the pos¬
terior margin.

The carpus of the first pereiopod is about twice as long as high ; it is
deeply excavated anteriorly. It is about as long as the merus, and shorter
than the chela. The chela is robust ; the fingers are somewhat longer than
the palm. The second legs are more slender. The carpus is more than 5 times
as long as high ; it is longer than either merus or chela. The fingers of the

Source : MNHN, Paris

14

1. HOLTHUIS

chela are slightly more than 1.5 times as long as the palm. The dactylus
of the third pereiopod ends in a dark claw, its posterior margin bears 5 to 7
additional teeth, the distal of which is distinctly smaller than either the
tooth that précédés it or the final claw ; from the second to the proximal.

Source : MNHN, Paris

ATYIDAE OF MADAGASCAR

15

the posterior teeth diminish regularly in size. A row of hairs is implan ted
on the latéral surface of the dactylus at the base of the distal two teeth.
The propodus is about 5 times as long as the dactylus and bears a row of
about 10 to 15 spinules on the posterior margin. The carpus is 0.7 to 0.8
times as long as the propodus. It bears a strong movable spine in the distal
part of the outer surface ; a few small spinules may be observed more proxi-
mally. The merus is about twice as long as the carpus ; it bears 3 or 4 strong
spines on the outer surface near the posterior margin. The isclüum bears
no movable spine. The dactylus of the fifth pereiopod ends in a curved
claw and bears numerous comb-like arranged slender spinules on the poste¬
rior margin. The propodus is 4.5 to 5 times as long as the dactylus ; its
posterior margin bears some 15 to 17 spinules, the distal of which are the
most conspicuous. The carpus is slightly more than half as long as the
propodus and bears a strong distal spine on the outer surface. The merus
is somewhat shorter than the propodus and is provided with 3 strong spines
in the distal half of the outer surface near the posterior margin.

The endopod of the first pleopod of the male is about triangular in outline
and possesses a well developed appendix interna, which reaches beyond
the top of the endopod. In the second pleopod of the male the appendix
masculina is about twice as long as the appendix interna ; it bears several
strong spines on its inner and distal margins. The diaeresis of the uropodal
exopod bears 19 to 23 spinules.

So far Caridina japonica was known only from Japan (from Sagami
Bay south) and Okinawa. The présent find of the species in four different
localities in Madagascar therefore cornes as a surprise. A comparison of the
Madagascar material with the good description and figures given by Kubo
(1938, p. 89, figs. 17-19) of Japanese specimens failed to produce any diffé¬
rence that might be considered of spécifie or subspecific value. Unfortu-
nately no Japanese material of the species is présent in the Leiden Muséum.

The species is closest, among Madagascar représentatives of the genus,
to Caridina iypus, from which it may at once be distinguished by the spinu-
lation of the dorsal margin of the rostrum. Also the shape of the dactyli
of the third and fourth periopods is quite characteristic.

Caridina nilotica (P. Roux) (fig. 5)

Pelias niloticus P. Roux, 1833, p. 73, pl. 7, fig. 1.

Caridina Wycki paucipara Bouvier, 1904, p. 130; Bouvier, 1905, p. 79.
Caridina wyckii Lenz, 1905, p. 385.

Caridina nilotica gracilipes Lenz, 1910, p. 568.

Caridina Voeltzkowi Lenz, 1910, p. 569.

Caridina nilotica typica Bouvier, 1925, p. 146 ; J. Roux, 1929,])]». 298, 318.
Caridina nilolica paucipara Bouvier, 1925, p. 153, figs. 316-318 ; J. Roux,
1929, pp. 300, 318.

Caridina nilotica Holthuis, 1956, p. 64.

Madagascar Institute :

— Ambila, Zanavorony Lake, about 150 km S of Tongobory, SW Madagascar ;
Aprll 1951 ; R. Paulian. — 1 spécimen.

Source : MNHN, Paris

, B. HOLTHUIS

— Ankavandra, Manambolo River, W Madagascar ; July 1949 ; R. Paulian. —
4 spécimens.

— N'amoroka, WSW of Kinkony Lake ; rivuiet ; September 1952 ; R. Paulian. —
12 specimens.

— Ambohiniorija Pool, Namoroka ; September 1952 ; R. Paulian. — 7 specimens

— Ambovonomby cave, Namoroka ; September 1952 ; R. Paulian. — 1 specimen

— Ampijoroa, Tsaramandroso, 30 km SE of Marovoay. — 4 specimens (1 ovigerous)

— Marovoay plain ; October 1949 ; R. Paulian. — li specimens (2 ovigerous).

— Antsohihy ; October 1949 ; R. Paulian. —11 specimens (4 ovigerous).

— Beangony at the foot of Tsaratanaro Mt., NE of Maromandia : October 1949
R. Paulian. — 5 specimens.

— Sambirano, Mizdioko, near Ambanja opposite Nosy Bé ; July 1948 ; J. Millot

— 3 specimens.

— Mananjeba River, Ambilobé, N Madagascar ; April 1951 ; R. Paulian. — 32 spe
cimens (3 ovigerous).

— Ambodivoangy, near Maroantsetra, Antongil Bay, NE Madagascar ; fresh water
1946 ; J. Millot. — I specimens (1 ovigerous).

— Alaotra Lake ; June 1951 ; P. A. Robinson. — 7 specimens.

— Highway from Ambalarondro to Anjahamanana near Brickaville, E Madagascar
September 1954 ; P. A. Robinson. — 7 specimens (1 ovigerous).

— Ranomafana near highway to Tamatave ; March 1954 ; P. A. Robinson. — 2 spe¬
cimens (1 ovigerous).

— Fort-Dauphin, SE Madagascar ; August 1948 ; J. Millot. — 13 specimens (1 ovi¬
gerous).

— Bevia Forest, Behara, S of Tranomaro, SE Madagascar ; August 1948 ; J. Millot.

— 8 specimens.

— Androy, Tranomaro; 11-15 August 1948; R. Paulian. — 2 specimens.

Austrian Madagascar Expédition :

— Lily River, an eftluent from Itasy Lake, W of Tananarive ; sandy shore ; 18 Sep¬
tember 1958 ; F.M. 228. — 6 specimens.

— Kinkony Lake, SW of Majunga, near west coast ; 19 June 1958 ; F.M. 246. _

1 specimen.

— Shore of Kinkony Lake ; 19 June 1959 ; F.M. 245. — 9 specimens.

— Bcmanondro Rivuiet, Nosy Bé ; 15 August 1958 ; F.M. 143. — 1 specimen.

— Andranobe Rivuiet, Nosy Bé ; 17 August 1958 ; F.M. 149. — 33 specimens.

— Fascène Rivuiet, Nosy Bé ; 18 August 1958 ; F.M. 154. — 7 juvéniles.

— Djabalabe Rivuiet, Nosy Bé ; 20 August 1958 ; F.M. 163. — 2 specimens.

— Djabala Rivuiet. Nosy Bé ; 22 August 1958 ; F.M. 172. — 5 juvéniles.

— Isaka Rivuiet, about 35 km NW of Fort-Dauphin, SE Madagascar ; 9 September
1958 ; F.M. 210. — 3 specimens (1 ovigerous).

— Rivuiet emptying in Anosy Lake, between Amboasary and Fort-Dauphin •
7 September 1958 ; F.M. 200. — 4 specimens (1 ovigerous).

Muséum Leiden :

— Rivuiet in Mitsinjo forest, Mitsinjo district, Majunga province, NW Madagas¬
car ; 25 September 1960 ; native naine * tsitsika * ; Y. Therezien no. 23. — 4 specimens/

— Northern Madagascar ; coll. Waterlot, don. Zoological Muséum Munich. — 5 spe¬
cimens (1 ovigerous).

— Antsakoamanondro rivuiet 13 km N of Ambanja on the highway to Ambilobé
Ambanja district, Diégo-Suarez province, N. Madagascar; 4 May 1959; Y. Therezien
no. 39, récolte no. 111. — 4 specimens.

— Beamalona River, Diégo-Suarez district, Diégo-Suarez province ; 28 April 1959 •
Y. Therezien no. 41, récolte no. 106. — 1 specimen.

— River at km. 107 of the highway from Diégo-Suarez to Ambilobé, Diégo-Suarez

district, Diégo-Suarez province ; 29 April 1959 ; Y. Therezien no. 42, récolte no. 108._

2 specimens.

— Andrarony River, Antalaha district, Diégo-Suarez province ; 8 km from the mouth
of the river; 7 February 1962; native naine • patsa ■ or « pitsikorana » ; Y. Therezien
no. 32. — 2 ovigerous fêmales.

— Lily-Ampefy-Itasy, Soavinandriana district, Tananarive province, Central Mada¬
gascar ; 18 September 1958 ; Y. Therezien no. 46, récolte no. 51. — 1 damaged specimen

— River near Anony Lake, between Amboasary du Sud and Fort-Dauphin, Amboa-
sary du Sud district, Tiiléar province, SE Madagascar; 7 September 1958; Y. Therezien
no. 37, récolte no. 35. — 2 specimens.

The specimens measure 14 to 30 mm. The ovigerous females are 17 to
30 mm long.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

17

The rostrum is very long, slender and curved upward at the tip. It
usually reaches distinctly beyond the antennal scale and has the tip bi-
or trifid. There is a distal unarmed portion on the dorsal margin behind
the single or double subapical tooth. The proximal part of the upper border
of the rostrum bears 13 to 27 teeth 2 or 3 of which are placed behind the orbit.
AU teeth are placed quite close together. The lower margin bears 6 to 22 teeth.
The upper teeth articulate with the rostrum, the lower do not. The lower
orbital angle is very distinct and separated from the antennal spine by a
good distance. The pterygostomian angle is rounded.

The dorsal surface of the telson bears 2 to 5, usually 3 or 4 pairs of
spinules. The posterior margin of the telson is rounded and shows no
médian tooth. The outer pair of posterior spines is very short, being less
than 1/4 of the length of the well developed latéral pair. There are 4 to
8 intermediate spines, which are undivided and distinctly shorter than
the laterals.

The stylocerite is slender and pointed, it reaches beyond the middle of
the basal segment of the antennular peduncle but distinctly fails to reach
the end of the segment. The antérolatéral tooth of this segment is sharply
pointed, it fails to reach the middle of the second segment. The spine near
the base of the scaphocerite is well developed and sharp.

The branchial formula is normal ; epipods are présent at the bases of
the first 4 pereiopods.

The carpus of the first leg is longer than the merus ; it is usually more,
sometimes somewhat less, than twice as long as liigh. It is shorter than the
chela. The carpus of the second leg is longer than the chela, it is 3 to 6 times
as long as high. In the third leg the dactylus bears 5 or 6 posterior teeth
additional to the distal claw ; the propodus is 4 to 4.5 times as long as the
dactylus. The propodus of the fifth legs is 3.5 to 4.5 times as long as the
dactylus.

The endopod of the first pleopod of the male is provided with a well
developed appendix interna. The exopod of the uropod bears 7 to 13 spinules
on the diaeresis. The preanal carina is blunt without a spine.

The eggs are rather voluminous, they measure 0.5-0.7 x 0.7-1.2 mm.

Caridina nilolica, if ail records in the literature are correct, is a very
widely distributed species, its range extending from eastern Africa (Egypte
to S. Africa) to Japan and Polynesia. A thorough study of the nilotica
complex is highly necessary and until then nothing positive about the area
of the species can be said. Type locality is the Nile river near Cairo. The
records from Madagascar found in the literature are the following : Mada¬
gascar (Bouvier, 1904, 1905, 1925), tributary of the Onilahy River near
Kofotsy at 50 km from the coast, SW Madagascar (Bouvier, 1904, 1925,
name of the river erroneously spelled Anilabry), Itasy Lake, Central West
Madagascar (Lenz, 1910 ; Bouvier, 1925), Tananarive (Bouvier, 1904,1925),
inundation swamp of the Betsiboka River (J. Roux, 1929), Majunga (Lenz,
1905), Namoroka and Kapiloza Rivers, Ambongo, Majunga province
(J. Roux, 1929), Ambovonomby Cave near Namoroka, Majunga province
(Holthuis, 1956), neighbourhood of Manarasandry, Majunga province
(J. Roux, 1929), Alaotra Lake, E Madagascar (Lenz, 1910 ; Bouvier, 1925,
who gave the locality under its correct name and under the incorrect spel-
lings Alasha, Maotra and Maotva), Imerimandrosa on Alaotra Lake (Bou-

Mémoires du Muséum. — Zoologie, t. XXXIII. 2

Source : MNHN, Paris

18

L. B. HOLTHUIS

Fio. 5. — Caridina nilolica (P. Roux), a, b, anterior part of body in latéral view ; c, tip
of telson ; d, pre-anal carina ; e, /, first pereiopod ; g, h, second pereiopod ; i, j, third
pereiopod ; k, /, fifth pereiopod ; m, endopod of flrst pleopod of male ; n, outer part
of diacresis. a, e-l, x 15 ; b-d, n, x 30 ; m, x 50. a, c-e, g, i, k, m, n, spccimen from
Mananjeba River ; b, specimen from Lily River ; /, h, /, I, specimen from Andranobe
rivulet.

vier, 1925 ; J. Roux, 1929), Ambatonharanana near Ampasimpotsy,
mountains leading to Mangoro Terrace (Bouvier, 1925, locality spelled
incorrectly Ambatoubavara), Ankobo, Manambovo River, N of Tamatave
(Bouvier, 1925, name of river spelled incorrectly Manambowo), Canal des
Pangalanes, région of Ambila, between Tamatave and Brickaville (J. Roux,

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

1929), Andevoranto, Lagoan district, S of Tamatave (Bouvier, 1925),
Vinanybe Lake, Fort-Dauphin, SE Madagascar (Bouvier, 1904,1925, name
erroneously spelled as la Vinangbe), Imanombo River, S Madagascar (Bou¬
vier, 1904) ; the localities Ivololo River (Bouvier, 1904, 1925) and Irangy
River (J. Roux, 1929) are unknown to me, while also Dr. Paulian, who
provided me with the exact position of many of the Madagascar localities,
informed me that he was unable to locate them.

Bouvier (1925) in his very useful monograph of the Atyidae recognized
a large number of “varieties” of Caridina nilotica. As shown already by
various authors, some these varieties are good species, while others cannot
be kept separate. As Gordon (1930) showed, part of Bouvier’s specimens
of Caridina nilotica var. typica do not belong to C. nilotica at ail, but are
C. africana Kingsley. Bouvier (1925) and J. Roux (1929) mentioned the
following varieties of C. nilotica for Madagascar : var. typica, var. styli-
roslris, var. xiphias, var. gracilipes, var. paucipara, var. brachydactyla,
and var. brevidactyla. After studying the présent material I hâve corne to
the conclusion that vars. xiphias and stylirostris are no C. nilotica and even
belong in a different section of the genus (see under C. xiphias). The speci¬
mens brought by Bouvier and J. Roux to vars. brachydactyla and brevi¬
dactyla I consider as specifically distinct from C. nilotica and |have assigned
them here more or less provisionally to C. longirostris. Part of the specimens
assigned by previous authors to var. gracilipes seem also to belong to
C. longirostris while others, judging by the description, might be what
I hâve named here C. nilotica. As to vars. typica and paucipara, I found
it impossible to distinguish between these two in my Madagascar material.
Bouvier (1925, p. 153) already remarked that the var. paucipara «passe
insensiblement à la forme typica, de sorte qu’il est souvent très difficile,
sinon impossible, d’attribuer certains individus à l’une ou l’autre des deux
variétés », an opinion which was fully shared by J. Roux (1929, p. 300).
Without giving any definite opinion about the status of the true Caridina
nilotica paucipara De Man from Natal, basing myself on the Madagascar
material I can only corne to the conclusion that we hâve to do here with
a single species of which I am unable to distinguish any subdivisions.

The status of Caridina nilotica and its various forms needs a careful
révision based on extensive collections from ail over the range of the species.
The fact that Bouvier’s (1925) species is a heterogeneous assemblage is
already clear from the fact that it is mentioned in no less than 6 places in
his key to the species of Caridina. A most useful contribution to our know¬
ledge of the status of the typical Caridina nilotica was given by Gordon
(1930) who examined material from the type area and compared it with
specimens from farther south in eastern Africa. Gordon’s material agréés
quite well with ours except for two characters, namely (1) that of the poste-
rior margin of the telson which in Gordon’s specimens is relatively narrower
and ends in a médian tooth (except for 4 aberrant specimens), while (2)
the eggs in her material are slightly smaller being 0.40-0.52 x 0.66-0.78 mm
in Egyptian specimens, and 0.31-0.45 X 0.54-0.73 mm in other specimens
from the Nile valley. As the character of the rounded tip of the telson is
not altogether constant in my material, while the différence in the size of
the eggs is so small, I thought it best to assign my material, for the time
being at least, to the typical Caridina nilotica. An examination of more

Source : MNHN, Paris

20

L. B. HOLTHUIS

material may prove me wrong here. The number of intermediate spines on
the posterior margin of the telson of Madagascar specimens agréés best
with that found by Gordon in material from the Southern part of the East
African range of the species.

Bouvier (1904, 1905, 1925) gave rather few details of the Madagascar
specimens which he assigned to C. nilotica lypica and C. n. paucipara. The
dimensions of the eggs is given by him (Bouvier, 1905, p. 80) as 0.5 x
0.65-0.9 mm, which agréés very well with what I found in my material.
Part of the specimens brought by Bouvier in 1904 and 1905 to C. n. pauci¬
para are referred by him to C. n. typica in 1925. In his 1925 book he did
figure a specimen (C. n. paucipara) from Madagascar (fig. 316-318).

Lenz (1905) gave too few details of his specimens to make certain
whether they belong to the présent or to the next species. The specimens
which the same author (Lenz 1910, p. 568, 569) assigned to C. n. gracilipes
and C. Voeltzkowi were identified by J. Roux (1929, p. 312), who evidently
examined them, as C. n. paucipara.

Caridina longirostris H. Milne Edwards (fig. 6)

Caridina longirostris H. Milne Edwards, 1837, p. 363.

Caridina Wyckii gracilipes Coutière, 1900, p. 1267.

Caridina nilotica brachydactyla Lenz, 1910, p. 568; Bouvier, 1925, p. 155.
Caridina nilolica gracilipes p.p. Bouvier, 1925, p. 152.

Caridina nilolica brevidadyla J. Roux, 1929, p. 303.

Madagascar Institute :

— St. Augustin, SW Madagascar; 4 April 1953; P. A. Robinson.

(1 ovigcrous.)

Agricultural Station Bas-Mangoky on the Southern shore of the Mangoky River
at 60 km from the sea ; March 1955 ; P. A. Robinson. — 3 specimens (2 ovigerous).

— River N of Majunga ; December 1951 ; R. Paulian. — 1 specimen.

Antsohihy, NW Madagascar; stagnant water ; October 1919; R. Paulian._

3 specimens.

— Ranomnfana near highway to Tamatave ; Mardi 1954 ; P. A. Robinson. — 4 spe¬
cimens (3 ovigerous). 1

specimens

Austrian Madagascar Expédition :

Djabala Rivulet, Nosy Bé, NW Madagascar ; 22 August 1958 ; F. M. 171. _

2 specimens (1 ovigerous).

— Androadroatra Rivulet, Nosy Bé ; 23 August 1958 ; F.M. 175. — 20 specimens

— Small lake near Mangatsa, Majunga district, NW Madagascar; 16 Januarv 1959 •
F.M. 240. — 2 specimens (I ovigerous).

— Mandromodromotra Rivulet near Mahialambo, about 20 km N of Fort-Dauphin •
11 September 1958; F.M. 217. — 5 specimens.

Muséum Leiden :

— Ampapamena near Analalava, Majunga province ; Waterlot coll. ; don. Muséum
Munich. — 2 specimens.

— Analalava, small brook near its mouth ; Waterlot coll. ; don. Muséum Munich
— 10 specimens (3 ovigerous).

— Antsirabe near Anorotsangana, Amboanga district, Nosy province ; Waterlot
coll. ; don. Muséum Munich. — 5 specimens (1 ovigerous).

- Ambatonjanahary near Diégo-Suarez, N Madagascar ; brook ; Waterlot coll. •
don. Mus. Munich. — 2 specimens.

— Near Ambahoabé, Simianona River. N of Soanierana, E Madagascar ; December
1938 ; H. J. Lam & A.D.J. Meeuse. — 1 ovigerous female.

Source : MNHN, Paris

THE ATYIDAE

.DAGASCAR

21

— Maningory, Tamatave province, E Madagascar ; 1927 ; G. Petit ; don. Mus. Munich.
— 1 specitnen.

— Mandromodromotra Rivulet, Piste Elandy, Fort-Dauphin district, Tuléar pro¬
vince, SE Madagascar ; 11 Scptember 1958 ; Y. Therezien no. 36, récolte no. 46. — 1 spé¬
cimen.

The specimens are 13 to 35 mm long, the ovigerous females measure
25 to 35 mm.

The rostrum usually is very long and slender, reaching distinctly beyond
the scaphocerite ; in a few specimens it is slightly shorter than the scapho-
cerite. The tip is curved upward, and bifid by the presence of a subapical
tooth. There is a distinct unarmed portion in the distal part of the upper
margin of the rostrum behind the subapical tooth. The upper border bears
13 to 23 proximal teeth, 2 or 3 of which are placed behind the orbit. The
lower margin bears 5 to 22 teeth. The lower orbital angle is well developed,
and distinctly separated from the strong antennal spine. The pterygostomian
angle is rounded.

The upper surface of the telson bears 2 to 4, usually 3 or 4, pairs of
spinules. The posterior margin of the telson is rather narrow and ends
in a sharply pointed médian tooth. The outer spines of this margin are
less than 1/3 of the length of the laterals. There are 2 to 6 intermediate
spines, which are not divided and are shorter than the laterals.

The eyes, antennulae and antennae show no différence from those of
C. nilotica.

The branchial formula is as in the previous species.

The carpus of the first pereiopod is longer than the merus and 2 to 3
times as long as wide. The carpus of the second pereiopod is more than
5 times as long as wide. The propodus of the third leg is 5 to 6 times as long
as the dactylus, that of the fifth leg measures 4.5 to more than 5 times the
length of the dactylus.

The endopod of the first pleopod of the male has a well developed
appendix interna, which is already quite distinct in juvénile specimens.
The exopod of the uropod shows 8 to 15 spinules on the diaeresis.

The preanal carina ends in a strong posteriorly directed sharp spine.

The eggs are numerous and small and measure 0.2-0.25x0.3-0.4 mm.

The Madagascar material brought to the présent species shows a very
close resemblance to C. nilotica but differs in a number of characters, the
most important of which are (1) the eggs are much smaller and more nume¬
rous, (2) the preanal carina bears a strong spine, (3) the posterior margin
of the telson ends in a médian point, (4) the dactyli of the last three pereio-
pods are relatively shorter.

Though I am convinced that the two forms indicated here as C. nilotica
and C. longirostris are distinct species, I am by no means certain that the
names given here are used correctly. Judging by Bouvier’s (1925) descrip¬
tion of the types of Caridina longirostris the présent form agréés closely
with it and for that reason it is provisionally assigned to this, one of the
longest known species of the genus. It will be up to future investigations
to define the status of the various species of Caridina, including that of
the présent one.

Coutière (1900) did not give any details of the Madagascar specimens
which he identified as Caridina wyckii gracilipes, but Bouvier (1925, p. 153)

Source : MNHN, Paris

22

i. HOLTHUIS

Fio. 6. — Caridina longirostris H. Milne Edwards, a, anterior part of body in latéral
view ; b, tip of telson ; c, pre-anal carina ; d, e, ilrst pereiopod ; /, g, second pereiopod ;
h i, third pereiopod ; J, k, fifth pereiopod ; l, m, endopod of iirst pereiopod of male ;
n ’outer part of diaeresis. a, d-k, x 13 ; b, x 26 ; c, x 52 ; /, m, x 40. a-d, /, h,/,
specimen from Ranomafana ; e, g, i, k, specimen from Analalava ; /, m, specimen
from Bas-Mangoky.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

23

under C. nilolica gracilipes mentioned « quelques spécimens portant l’indi¬
cation H.C. 19 (3 paires d’épines dorsales au telson, œufs de 0.40) », which
may hâve been Coutière’s specimens, and which, judging by the size of
the eggs, belong in the présent species.

Lenz's (1910, p. 569) specimens indicated with the name Caridina
nilotica brachydaclyla evidently also belong here in view of the fact that
one of the characters of de Man’s form is that the dactyli of the last three
pereiopods are short. In my opinion Caridina brachydaclyla is a distinct
species, characterized among other things by the strong development of
the distal spine on the posterior margin of the last three pereiopods. No
Madagascar specimens of that species hâve been seen by me, the Mada¬
gascar specimens assigned to that form by previous authors in ail proba-
bility ail belong to C. longiroslris. J. Roux’s (1929) account of his Madagascar
C. nilolica brev dactyla also checks well with our specimens. Therefore ail
are provisionally identified with Caridina longirostris.

Due to the confusion existing in the systematics of the nilolica group
of Caridina it is impossible to give the exact range of the présent species.
It was originally described from Algeria, but authors like Bouvier (1925,
p. 149-151) hâve made it very probably that this locality indication is
erroneous. It is possible that the species has a rather wide distribution in
the Indo-West Pacific area. The previous Madagascar records are the
following : Madagascar (Coutière, 1900), Onilahy River, SW Madagascar
(Bouvier, 1925), Maroamalona, Tuléar Province (Bouvier, 1925), Kinkony
Lake, W Madagascar (Lenz, 1910, spelled as Kikuni), Ivoloina River near
Tamatave, E Madagascar (J. Roux, 1929), and the locality Andamagassa
(Bouvier, 1925), the situation of which is unknown to Dr. Paulian and
myself, and probably is incorrectly spelled.

Caridina gracilirostris De Man (fig. 7)

Caridina gracilirostris De Man, 1892, p. 399, pl. 25, fig. 31 ; J. Roux, 1929,

p. 296.

Madagascar Institute :

— Agricultural Slation Bas-Mangoky on the Southern shore of the Mangoky River
at 60 km from the sca, SW Madagascar ; March 1955 ; P.A. Robinson. — 12 specimens
(4 ovigerous).

— Marovoay plains, NW Madagascar ; October 1949 ; R. Paulian. — 3 specimens
(2 ovigerous).

Muséum Leiden :

— Antsirabe near Anorotsangana, Amboanga district, Nosy province, NW Mada¬
gascar ; Waterlot coll., Muséum Munich don. — 3 specimens.

— Ambanja Lake, Ambato-Boeni district, Majunga province, NW Madagascar ;
3 November 1960 ; native names ■ tsitsika » and • tsivaky » ; Y. Thcrezien nos. 27 and 30.
— 21 specimens (6 ovigerous).

The specimens measure 21 to 37 mm ; the ovigerous females are 26 to
35 mm long.

The rostrum is very long and slender, reaching distinctly beyond the
scaphocerite, often with more than half its length. The tip of the rostrum

Source : MNHN, Paris

24

L. B. HOLTHUIS

is curved upwards, it is bifid to quadrifid by the presence of one to three
subapical teeth. A considérable portion of the dorsal margin behind the
subapical tooth is entire. In the proximal part the dorsal margin bears 5
to 13 teeth of which at most 1 is placed behind the orbit. The teeth are
rather widely separated. The lower margin of the rostrum is serrate and
bears 28 to 33 teeth. The lower orbital angle is very distinct and rounded.
The antennal spine is strong, and sharply pointed ; it is placed some distance
below the orbital angle. The pterygostomian angle is not produced but
about rectangularly rounded.

The telson bears dorsally 3 or 4 pairs of spinules. Its posterior margin
is rather narrow and ends in a distinct médian point. The outer spines are
about half as long as the laterals. The intermediate spines are absent or
represented by a single pair.

The stylocerite is slender and pointed, it reaches beyond the middle of
the basal segment of the antennular peduncle but fails (sometimes by not
very much) to attain the end. The antérolatéral tooth of the segment is
well developed but fails by far to reach the middle of the second segment
of the peduncle. At the base of the scaphocerite the antennal peduncle
shows a well developed sharp spine.

Epipodites are présent at the base of the first four pereiopods.

The carpus of the first cheliped is longer than the merus and about 2
to 2.5 times as long as wide. The carpus of the second leg is more than 5

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

25

times as long as wide. In the third pereiopod the propodus is more than
4 times as long as the dactylus, while in the fifth pereiopod the propodus
is 3.5 to 4 times the length of the dactylus.

The endopod of the first pleopod of the male bears a distinct appendix
interna. The diaeresis of the uropodal exopod bears 8 to 12 spinules. The
preanal carina ends in a distinct posteriorly directed spine.

The eggs measure 0.3-0.4 x 0.5-0.6 mm.

In many respects the présent species is extremely close to C. longirostris,
it may immediately be recognized however by the small number of dorsal
teeth of the rostrum, which are very widely spaced. This feature is so striking
that it serves as an easy means to identify the species.

The présent specimens agréé well with the descriptions given of the
species in the literature.

The species is known from a large part of the Indo-West Pacific area
(Madagascar and India to Malaysia and Palau). The only previous record
of the species from Madagascar is that by J. Roux (1929) who reported
it from the Maintimazo River at Bemazaka, Majunga province, N.W. Mada¬
gascar.

Caridina serratirostris De Man ( fig. 8)

Caridina serratirostris De Man, 1892, p. 382, pl. 23, fig. 28.

Madagascar Instituts :

— Antsohihy, NVV Madagascar; stagnant water ; October 1949; R. Paulian. —
1 specimen.

Austrian Madagascar Expédition :

— Small Iake near Mangatsa, Majunga district, NW Madagascar ; 16 January 1953;
F.M. 240. — 3 specimens.

Muséum Leiden :

— Antsirabe near Anorotsangana, Amboanga district, Nosy province, NW Mada¬
gascar ; leg. M. Waterlot ; don. Mus. Munich. — 2 specimens (1 ovigerous).

The specimens are 9 to 20 mm long, the ovigerous female measuring
20 mm.

The rostrum is straight, it reaches slightly beyond the end of the anten-
nular peduncle. The upper margin is evenly serrated bearing 19 to 26 teeth,
6 to 8 of which are placed behind the orbit. The lower margin bears 6 to 8
teeth, the last being removed some distance from the tip of the rostrum.

The antennal spine is distinct, it is placed some distance below therounded
lower orbital angle. The pterygostomian angle is rounded, not produced.

The telson bears 4 or 5 pairs of spines on the dorsal surface. The posterior
margin is rounded with a subterminal médian spinule. The latéral spines
of the posterior margin are more than 4 times longer than the outer. The
outer pair of intermediate spines is shorter than the latéral and simple,
between these there are 4 (in juvéniles) to 8 (in old specimens) long spines,
which are distinctly longer than the laterals and are divided in two by a
kind of chitinous plug.

Source : MNHN, Paris

26

L. B. HOLTHUIS

The eyes are well developed with a rounded pigmented cornea. The
stylocerite is very long, reaching far beyond the antérolatéral angle of the
basal segment of the antennular peduncle and reaching to or even beyond
the middle of the second segment. This second segment is exceptionally
long and slender. The stylocerite is relatively longer in adults than in juvé¬
niles but always reaches beyond the basal segment. The scaphocerite ends
is a strong tooth, which is hardly overreached by the lamella. There is a
strong and slender outer spine on the antennal peduncle near the base
of the scaphocerite.

Fio. 8. — Caridina serratirostris De Man. a, anterior part of body in latéral view
b, eye, antennula and anlenna, in dorsal view ; c, tip of telson ; d, third maxilliped
e, flrst pcreiopod ; /, second pereiopod ; g, third pereiopod ; h, flfth pcreiopod ; i, en
dopod of flrst pleopod of male ; j, outer part of diaeresis. a, b, d-h, x 16 ; c, /, x 32
i. x 65. a-h, j, specimen front Antsirabe ; i, specimen from Antsohihy.

Source : MNHN, Paris

THE ATYIDAE

MADAGASCAR

27

Epipods are présent on the bases of the first four pereiopods.

The carpus of the first pereiopod is longer than the merus, being 3.5 times
as long as wide and about as long as the chela. In the second pereiopod the
carpus is far longer than the chela and about 10 times as long as wide. The
fingers are remarkably long and slender. The dactylus of the third pereiopod
is relatively very slender, it ends in two teeth and bears about 9 additional
spinules on the posterior margin. The propodus is about 3 J times as long as
the dactylus. The dactylus of the fifth leg is also very slender, bearing
about 16 spinules on the posterior margin behind the distal pair. Here too
the propodus is about 3.5 times as long as the dactylus.

The preanal carina ends in a strong posteriorly directed spine.

The uropodal exopod bears 15 to 18 spinules on the diaeresis.

The eggs are numerous and small, measuring 0.3-0.35 X 0.5 mm.

The species is very characteristic and immediately recognisable by the
large number of dorsal rostral teeth behind the orbit and by the long
stylocerite.

It was originally described from the Lesser Sunda Islands, Indonesia,
and is now known to inhabit an area extending from the Seychelles and
Mauritius to Okinawa, Polynesia, and N.E. Australia. It is herereported
for the first time from Madagascar.

Caridina petiti J. Roux

Caridina petiti J. Roux, 1929, p. 314, figs. 13-15 ; Decary, 1950, p. 167.

No material of this species has been examined by me. So far it is only
known from the type material. For the description I refer to J. Roux’s
(1929) account.

The species shows closest resemblance to C. calmani with which it agréés
in most respects. The most conspicuous différence being the full dentition
of the rostrum in C. petiti. It is possible that C. petiti should be considered
to be a subspecies of C. calmani.

The type-locality of J. Roux’s species is eastern lagoons near Ambila
(between Brickaville and Tamatave), eastern Madagascar.

Caridina calmani Bouvier (fig. 9)

Caridina Calmani Bouvier, 1919, p. 334 ; Bouvier, 1925, p. 253, figs. 572-599.
Caridina Bouvieri J. Roux, 1929, p. 312, figs. 7-12.

Madagascar Instituts :

— Highway near Anosibé, SE of Tananarive, E Madagascar ; running water; January
1951 ; R. Pauîian. — 20 specimens (2 ovigerous).

— Foresl near Anosibé ; January 1951 ; R. Paulian. — 4 specimens (2 ovigerous).

— Near Forestry Station Manjakatompo, Ambahona, Ankaratra massif, above
Ambatolampy, 70 km S of Tananarive : brooklets and pools ; altitude 1700 m; 4 Juiy 19o6.
— 4 specimens.

Source : MNHN, Paris

28

L. B. HOLTHUIS

Austrian Madagascar Expédition :

— Périnet, E Madagascar (halfway between Tananarive and thc coast); small brook
emptying in Lac Vert ; 28 July 1958 ; F.M. 109. — 2 specimens (1 ovigerous).

— Périnet ; small brook running from Lac Rouge ; 23 July 1958; F.M. 111. — 5 spe¬
cimens.

— Périnet ; small brook ncar Lac Rouge ; 23 July 1958; F.M. 112. — 7 specimens.

— Périnet ; source in Virgin forest ; 28 July 1958 ; F.M. 113. — 32 specimens.

— Périnet ; source in swampy part of virgin forest ; 30 July 1958; F.M. 117. _

14 specimens.

— Périnet ; source ; 30 July 1958 ; F.M. 118. — 28 specimens (2 ovigerous).

— Périnet ; source in virgin forest ; 31 July 1958; F.M. 120. — 2 specimens (1 ovi¬
gerous).

The specimens are 6 to 20 mm long. The ovigerous females measure
14 to 16 mm.

The rostrum is short, it reaches to the base or to the end of the second
segment of the antennular peduncle, sometimes it is shorter than that,
rarely longer. As a rule the rostrum is rather high, but sometimes quite
shallow. The upper margin bears no teeth, rarely 1 dorsal tooth is found
and one specimen even showed 7 teeth. The lower margin bears 0 to 3 teeth.

The antennal spine is placed the lower below orbital angle. Sometimes
it is well developed and sharply pointed, sometimes it is just a blunt
angle. The lower orbital angle is usually quite inconspicuous and often folded
inwards and thereby obscured from view. The pterygostomian angle is
rectangularly or more acutely rounded, it is as a rule distinctly forwards
produced, sometimes very much so.

The dorsal surface of the telson bears 3 to 5 pairs of spinules. The poste-
rior margin of the telson is rounded without a médian point. The latéral
spines are more than 4 times longer than the outer. The intermediate spines
are 4 to 14 in number, the outer of them are shorter, the inner longer than
the latéral spines, they are divided in two by a chitin plug.

The eyes are well developed. The antennula is robust with short segments.
The stylocerite is rather blunt, reaching beyond the middle of the basal
segment, but failing to reach the end of it. The antérolatéral angle of the
segment is produced into a tooth, which does not attain the middle of the
second segment. The antennal peduncle shows a distinct tooth near the base
of the scaphocerite. This tooth may be broad and blunt or rather slender.
The final tooth of the scaphocerite is broad.

Epipods are found at the bases of the first four pereiopods.

The carpus of the first pereiopod is slender, it is longer than the merus
and about as long as the chela ; it is 2.5 to 3 times as long as wide. The
carpus of the second pereiopod is still much more elongate, being 6 to 7
times as long as wide and much longer than either merus or chela. The
propodus of the third leg is about 3 to 4 times as long as the dactylus. The
propodus of the fifth leg is also about 3 to 4 times as long as the dactylus.
The endopod of the first pleopod of the male is without appendix interna!

The preanal carina is blunt without a spine.

The diaeresis of the uropodal exopod bears 12 to 20 spinules.

The eggs are large, they measure 0.6-0.7 X 0.9-1.1 mm.

The species sofar is only known from central eastern Madagascar. The
type locality is Ambatonharanana near Ampasimpotsy, in mountains
leading to the Mangoro Terrace near lake Alaotra (erroneously indicated
by Bouvier as Ambatoubavara). The range seems to be rather restricted.

Source : MNHN, Paris

\yjL

THE ATYIDAE OF MADAGASC/

29

F ,g - 9- — Caridina calmant Bouvier, a-d, anterior part of body in latéral view : e, tip
of telson ; /, third maxilliped ; g, h, first pereiopod ; i, j, second pereiopod ; k, 1, third
pereiopod ; m, n, flfth pereiopod ; o, endopod of first pleopod of male : p, outer part
of diaeresis a-d /-n x 16 ; e, p, x 32 ; o, x 65. a, e-g, i, k, m, o, p, specimen fVom
Permet, F.M. 118 ; b, h, /, I, n, specimen from highway near Anosibé ; c, d, specimen
from Permet, F.M. 120. F

Source : MNHN, Paris

30

L. B. HOLTHUIS

Examination of topotypic material of Caridina bouvieri J. Roux con-
vinced me that that species cannot be separated from C. calmani, and
represents at most a local variant characterized i.a. by a low number of
spinules on the diaeresis.

Caridina angulata Bouvier

Caridina angulata Bouvier, 1905, pp. 75, 84, fig. 8 ; Lenz, 1910, p. 570 -
Bouvier, 1912, p. 919; Bouvier, 1913, p. 465; Bouvier, 1925, p. 240*
figs. 555-561.

I hâve seen no material of this species. It has been reported from an
area roughly SW of that of C. calmani. An examination of a more extensive
material from both areas may prove that the two species actually are only
subspecies ; in that case the name angulata has priority. The type locality
of C. angulata is Ranofotsy River near Fianarantsoa, at the plateaux
between Ihosy and Tananarive. Lenz (1910) reported the species from
Itasy Lake.

Caridina edulis Bouvier

Caridina edulis Bouvier, 1904, p. 135 ; Bouvier, 1905, p. 91, fig. 13 ; Bouvier

1912a, p. 919 ; Bouvier, 1913, p. 465 ; Bouvier, 1925, p. 208, figs. 458-463*

This species of which no material has been examined by me, seems to
be characterized by the dactyli of the third legs, which according to Bouvier
( 1925, p. 210, fig. 460) hâve 15 to 17 teeth on the posterior margin, a number
which is more than double that of the related Madagascar species.

The types are known to corne from Madagascar, but no more précisé
locality indications are available. Bouvier (1904) brought also some spéci¬
mens from Anantsahalankely to this species.

Caridina isaloensis isaloensis Coutière (fig. 10)

Caridina lypus isaloensis Coutière, 1899, p. 383.

Caridina typus Isaloensis Coutière, 1900, p. 1267.

Caridina isaloensis Bouvier, 1904, p. 135; Bouvier, 1905, p. 90, fig. 12 -
Bouvier, 1912a, p. 919 ; Bouvier, 1913, p. 465 ; Bouvier, 1925, p. 195’
figs. 412-415 ; J. Roux, 1929, p. 310 ; Decary, 1950, p. 167.

Caridina madagascariensis Bouvier, 1904, p. 134; Bouvier, 1905, p. 89-
Bouvier, 1912, p. 292; Bouvier, 1912a, p. 919; Bouvier, 1913, p. 464-
Bouvier, 1925, p. 200, figs. 428431 ; J. Roux, 1934, p. 530.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

31

Madagascar Instituts :

— Bekily, N of Imanombo, Onilahy River basin, S Madagascar ; August 1948 ;
J. Millot. — 7 specimens.

— Betroka, about 150 km S ol Ihosy, Onilahy River Basin ; July 1948; J. Millot. —
5 specimens.

— Bezaha on Onilahy River ; April 1953 ; P.A. Robinson. — 4 specimens.

— Sept Lacs, Tongobory, Onilahy River basin ; May 1951; R. Paulian. — 20 spe¬
cimens.

— Ankazoabo, SW Madagascar ; November 1936; P.A. Robinson. — 21 specimens.

— Ankaramena, Mangoly River basin, SW Madagascar; April 1955; E. Razaflman-
dimby. — 13 specimens.

Austhian Madagascar Expédition :

— Iritsoka brook, a few km N of Betroka, S of Ihosy, Onilahy River basin; 13 Sep-
tember 1958 ; F.M. 222. — 14 specimens (1 ovigerous).

— Brook near Ambohimanavelona, SE of Tuléar, on the highway to Tongobory,
Onilahy River basin, SW' Madagascar ; 4 Septcinber 1958 ; F.M. 190. — 25 specimens
(3 ovigerous).

Muséum Leiden :

— Iritsoka Brook near km 94 on the highway from Ihosy to Fort-Dauphin, 24 km N
of Betroka, Betroka district, Tuléar province, S Madagascar ; 13 September 1958 ; Y. The-
rezien no. 38, récolte no. 49. — 3 specimens.

The specimens are 7 to 24 mm long, the ovigerous females being 14 to
24 mm.

The rostrum in typical specimens reaches to the base or to the middle
of the second segment of the antennular peduncle, seldom the end of the
segment is reached. The tip is often slightly curved up. The upper margin
of the rostrum is typically unarmed ; a few (6 of 78) specimens in some of
the lots examined (from Iritsoka, Bekily, Betroka) showed some (1 to 7)
dorsal teeth, in the lot from Ankazoabo there were 9 specimens with (3 to 12)
dorsal teeth. The rostrum is high in some specimens, being much flatter
in others. The lower margin bears 0 to 2 seldom up to 4 teeth.

The lower orbital angle is not very distinct, generally it is folded inwards
so that it seems as if the antennal spine is placed on this lower angle. The
antennal spine is distinct and sharply pointed. The pterygostomian angle
is rounded and only slightly produced.

The dorsal surface of the telson bears 2 to 5, usually 3 pairs of spinules.
The posterior margin is rounded ; the outer spines are short and less than
1/4 of the length of the laterals, the intermediate spines are 3 to 9 in number
and are shorter than the laterals.

The eyes are well developed. The antennular peduncle is rather slender.
The stylocerite is sharp, reaches beyond the middle of the basal segment
of the antennular peduncle, but fails to reach the end. The antérolatéral
tooth of the segment is much shorter than half the second segment of the
peduncle. The antennal peduncle shows a usually distinct, often rather
broad and inconspicuous spine near the base of the scaphocerite.

Epipods are présent on the bases of the first four pereiopods.

The carpus of the first pereiopod is usually not very deeply excavated
anteriorly, it is 1.7 to 2.5 times as long as high, it is as long as the merus
and shorter than the chela. The carpus of the second leg is about 4 to 5.5
times as long as high ; it is longer than both merus and chela. The dactylus
of the third leg bears 4 to 6 rarely 7 teeth on the posterior margin. The pro-
podus is 3 to 4 times as long as the dactylus. Also in the fifth leg the propodus
is 3 to 4 times the length of the dactylus.

Source : MNHN, Paris

32

L. B. HOLTHUIS

The endopod of the first pleopod of the male is ovate and bears no
appendix interna.

The preanal carina is rounded without a tooth.

The diaeresis of the uropodal exopod bears 7 to 12, sometimes 13 spinules

The eggs are rather large, measuring 0.5-0.65 X 0.8-1.0 mm.

The présent species seems to hâve a rather wide distribution in Mada¬
gascar, but is represented by at least two local subspecies which graduallv
merge into each other.

The main characteristic of the présent, typical subspecies is the short
dorsally unarmed rostrum. In some populations there are specimens in
which the rostrum shows some dorsal spinules ; these specimens form a
transition to the next subspecies C. i. grandidieri Bouvier. The typical
subspecies seems to inhabit the SW corner of Madagascar.

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

33

Bouvier (1925) distinguished C. isaloensis from C. madagascariensis
and treated the two as distinct species. However, the différences mentioned
by Bouvier fall within the range of variation of the species and I cannot
find any good reason not to synonymize the two.

The species is only known from Madagascar. The type locality is the
Onilahy River in SW Madagascar (Coutière, 1899). Coutière (1900)
reported the species from "Madagascar (côte est)”, which clearly is an error
as the original description gives "(Côte Ouest)”. Bouvier (1904) reported
on about 40 specimens collected by Grandidier in 1898 between Salvamby
and Manasco (spelled later by Bouvier, 1925 : Saloambany and Manasca) ;
as the size of the lot, the collector and the approximate date are the same
as those of the type lot, I agréé with Bouvier (1925) that it probably is
the type lot. The other records of the typical subspecies are : Madagascar
(Bouvier, 1905, 1912, 1913), Mahafaly, Tranoroa région, S Madagascar
(J. Roux, 1934), Andranolaho, Onilahy River, Betsioky district (Bouvier,
1925 ; locality misspelled Andrandabo), Tuléar (J. Roux, 1934), Andrana-
kanga, Ambongo near Namoroka (J. Roux, 1929), Ankavandra (Bouvier,
1925), Fandramanona River (Bouvier, 1904 ; Bouvier, 1905 ; Bouvier,
1925 ; locality in 1925 misspelled Fandiamanana). The last mentioned loca¬
lity is not known to Dr. Paulian, it is possible that not only the second but
also the first spelling is incorrect ; this is the more to be regretted as the
locality is the type locality of both C. madagascariensis Bouvier and C. gran-
didieri Bouvier.

Caridina isaloensis grandidieri Bouvier (fig. 11)

Caridina Grandidieri Bouvier, 1904, p. 133 ; Bouvier, 1905, p. 87, fig. 9 ;
Bouvier, 1912, p. 293 ; Bouvier, 1912a, p. 919 ; Bouvier, 1913, p. 464 ;
Bouvier, 1925, p. 200, figs. 423-426.

Madagascar Institute :

— Ihosy, basin of Mangoky River ; July 1948 ; J. Millot. — 12 specimens.

— Canyon of Manambolo River, \V Madagascar ; pool in torrent near higliway ;
July 1949. — 7 specimens.

— Betsingily on Manambolo River : July 1949 ; R. Paulian. — 2 specimens.

— 23 km S of Ankavandra, Manambolo River ; pool in torrent ; R. Paulian. —
16 specimens.

— 20 km S of Ankavandra, Manambolo River ; pool in a torrent; under a piece of
wood ; 2 May 1949 ; P. Cachan. — 27 specimens.

— Bekopaka, S of Antsingy, Manambolo River basin ; July 1949; R. Paulian. —
36 specimens (3 ovigerous).

— N of Antsingy. — 13 specimens.

— Ambina, N of Antsingy ; July 1949: R. Paulian. — 16 specimens.

— Morafénobé, Mahajeby forest, W Madagascar ; May 1952; R. Paulian. — 15
specimens.

— Namoroka, WSW of Kinkony Lake, NW Madagascar ; in brook ; September 1952;
R. Paulian. — 4 specimens.

— Ampijoroa, Ankarafantsika, 30 km SE of Marovoay on the highway Majunga-
Tananarive ; ait. 120 m ; P. Grio; 4 September 1956. — 8 specimens (2 ovigerous).

— Maromandia, N Madagascar ; torrent ; July 194S; J. Millot. — 9 specimens.

— Ambilobé, N Madagascar ; Mananjeba River, April 1951; R. Paulian. — 3 spe¬
cimens (1 ovigerous).

Muséum Leiden :

— Ampombilava, Befandriana du Nord district, .Vlajunga province, N Madagascar;
24 June 1959 ; Y. Therezien no. 45, récolte no. 115. — 6 specimens.

Mémoires

Muséum. — Zoologie, t. XXXIII.

Source : MNHN, Paris

L. B. HOLTHUI

34

Fig. 11. — Caridina isaloensis grandidieri Bouvier, a-/, anterior part of body in latéral
view : g-i, ilrst pereiopod ; /-/, second pereiopod ; m-o, third pereiopod ; p-r, flfth

This subspecies is very closely related to the typical subspecies and the
only worth while différence that I could find is in the shape and dentition
of the rostrum. The rostrum is somewhat longer, reaching the middle or

Source : MNHN, Paris

THE ATYI

E OF MADAGASCAR

35

the end of the second segment of the antennular peduncle (though in some
specimens it only reaches the base of this segment and in very few it is
somewhat longer). The upper margin of the rostrum bears as a rule 6 to

18 teeth, seldom less (2 to 5) or more (19 to 24) ; of these 0 to 2 are placed
behind the orbit, sometimes the distal teeth are wider spaced than the proxi¬
mal, in a few cases the distalmost part of the upper margin is unarmed.
The teeth are short. The lower margin bears 0 to 10 teeth, which often are
very inconspicuous. The rest of carapace is as in the typical subspecies, as
are also the cephalic and thoracic appendages.

The diaeresis bears 8 to 13 (seldom 15) spinules, in the northernmost
lots, viz., those from Maromandia and Ambilobe these spinules are up to

19 in number.

This subspecies occupies a locality in W Madagascar north of that
from which the typical subspecies is known. Intermediate forms between
the two subspecies are found, and it is difficult to draw the exact line
between the two, it seems best, however, to keep these two forms separate.
The type locality of the présent subspecies is Fandiamanana River, the
exact position of which unfortunately is not known, it is possible that
Bouvier spelled the name wrong (the spelling Fandramanona River lias
also been used by Bouvier for this river). Bouvier (1904) doubtfully
referred specimens from Mavatana (situation unknown to me) to the présent
form. There are no other records known.

Caridina norvestica n. sp. (fig. 12)

Madagascar Institute :

— River N of Majunga ; December 1951 ; R. Paulian. — 6 specimens (1 ovigerous).

— Near Majunga, small brook at km 530 of the highway to Tananarive ; October
1949 ; R. Paulian. — 14 specimens.

— Maliajamba Lake ; October 1949 ; R. Paulian. — 6 specimens (2 ovigerous).

The specimens are 12 to 20 mm long ; the ovigerous females measure
19 to 20 mm.

In C. norvestica the rostrum is long, in the adults it reaches to or slightly
beyond the end of the antennular peduncle, and as a rule is curved upward
at the end. In juvéniles it may reach slightly beyond the base of the second
segment of the peduncle. One or two subapical teeth are présent, behind
which there is a conspicuous unarmed portion. In the basal part of the
rostrum there are 10 to 27 teeth ; in the adults the number of these teeth
(19 to 27) is much higher than in juvéniles (10 to 18). One or two of these
teeth are placed behind the posterior margin of the orbit. The lower margin
of the rostrum has 4 to 15 teeth, here too the number is higher in the larger
specimens than in the smaller.

The lower orbital angle is distinct and narrowly rounded. The well
developed antennal spine is placed some distance below the orbital angle.
The pterygostomian angle is rounded and only slightly produced.

The 6th abdominal somite is 0.6 to 0.7 times the length of the carapace.

The dorsal surface of the telson béais 3 or 4 pairs of spines. The posterior

Source : MNHN, Paris

L. B. HOI.THUIS

margin of the telson is rounded the outer spines are very short the lalerals
are more than 4 limes longer. The intermediate spines are 3 to 5 in number
and are almost as long as the laterals. The preanal carina is blunt.

The eycs are well developed and rather large. The antennal ridge is
somewhat elevated and rounded. The stylocerite is sharp and slender
reaching beyond the middle of the first antennular segment, but failing
to attain the end. The antérolatéral toolli of the segment is well developed
but fails to reach the middle of the quite slender second antennular segment.
The scaphocerite is slender, the spine near its base is broadly triangular
and inconspicuous.

Epipods are présent at the bases of the first four pereiopods. The carpus
of the first cheliped is 1.2 to 2.2 times as long as high, it is of about the
same length as the merus and shorter than the chela, it is not deeply excavate
anteriorly. The carpus of the second cheliped is about 5 to 5.5 times as long
as high ; it is distinctly longer than either chela or merus. The dactylus of

the third pereiopod bears 5 to 7 teetli on the posterior margin. The propodus
is about 3.7 to 4 times as long as the dactylus. The carpus bears one strone
movable spine in the distal part of its outer surface, the merus also possesses
such a spine and has moreover 2 more placed lower down near the posterior
margin of the segment. One such spine is présent on the ischium. The dactv-
lus of the fifth leg bears a row of numerous comb-like arranged spinules on

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

37

the posterior margin. The propodus is about 3.5 to 4.3 times as long as the
dactylus and bears several small posterior spinules. The carpus bears a
movable spine similar to that of the carpus of the third leg. In the merus
there are usually only 2 instead of 3 spines, wliile the ischium shows no
spine at ail.

The endopod of the first pleopod of the male is ovate without an appendix
interna. The appendix masculina of the second pleopod is somewhat longer
and broader than the appendix interna. The diaeresis of the exopod of the
uropod bears 9 to 15 spinules.

The eggs measure 0.5-0.7 x 0.8-1.0 mm.

The présent species is closely related to C. isaloensis and agréés with
that species in the general shape of the appendages, the telson and the
antennal spine. The most important différence being that of the rostruin,
which especially in adult specimens is greatly different. However, it may
ultimately prove necessary to reduce the présent species to the rank of a
subspecies of C. isaloensis.

Caridina troglophila n. sp. (fig. 13;

Caridina ? gladiifera Holthuis, 1956, p. 65.

Madagascar Institute :

— Ambovonomby cave, N'amoroka, NW Madagascar ; September 1952; R. Paulian.
— 32 specimens.

The specimens measure 14 to 25 mm.

The rostrum is rather high it reaches to or slightly beyond the end of
the antennular peduncle, it is straight or slightly curved up at the tip. The
upper margin is beset by 23 to 35 teeth, 4 to 6 of which are placed behind
the posterior orbital margin. The teeth are small, the distalmost are smallest,
often rather inconspicuous and more widely spaced than the proximals.
The lower margin bears 5 to 9 teeth ; these are low and not distinctly set
olT from the body of the rostrum, resembling thereby the distalmost dorsal
teeth.

The antennal spine is well developed and sharp, it is placed some distance
below the rounded orbital angle. The pterygostomian angle is broadly
rounded, and may be slightly produced.

The sixth abdominal somite is about 2/3 of the length of the carapace
(rostrum excluded). The telson bears dorsally 4 or 5 pairs of spines. The
posterior margin of the telson is not rounded but narrowed in three steps
resembling thereby the stepped gable of old dutch houses. The outer pair
of posterior spines is short, being less than 1/4 of the length of the long
latéral spines. The intermediate spines are 3 to 5 in number ; they are
shorter, but not very much shorter, than the latéral spines, and are simple,
not divided. The preanal carina is blunt, without a spine.

The eyes are well developed, but still relatively smaller than in related
species like C. noroestica. The cornea is distinctly pigmented and globular.
The antennular carina is rather distinct and somewhat produced. The

Source : MNHN, Paris

38

L. B. HOLTHUIS

stylocerite is slender and pointed, it reaches a little beyond the middle
of the basal segment of the antennular peduncle. The antero-lateral tooth
of the segment is slender and sharp but falls far short of the middle of the
second segment.

The scaphocerite reaches somewhat beyond the rostrum. The blade
distinctly overreaches the final tooth. There is a sharp, well developed spine
on the outside of the antennal peduncle near the base of the scaphocerite

The branchial formula is normal. The first four pereiopods each bear
an epipod at the base.

Fig. 13. — Caridina Iroglophila

part of body in latéral view ; b, tip of telson ; c, third maxillip.
e, second pereiopod ; /, third pereiopod ; g, fifth pereiopod
pleopod of male, a, c-g, x 10 ; b, i, x 32 ; h, x 50.

nc« species, from Ainbovonomby cave, a
b, tip of telson ; c, third maxilliped ; d, first pereiono,^
ifth nereionod : h. «ndopod of 1 first

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

39

The third maxilliped reaches as far as the final tooth of the scaphocerite.
Its distal segment is about as long as the penultimatc and bears spines
in the distal half of its lower margin. The first pereiopod is short and stubby.
The carpus is almost 1.5 times as long as high, it is deeply excavated ante-
riorly, it is longer tlian the merus but shorter than the chela. The Angers
are slightly shorter tlian the palm. The second legs are slender. The carpus
is 5 times as long as high, it is longer than both the merus and the chela.
The fingers are distinctly longer than the palm. The dactylus of the third
leg bears about 6 teeth on the posterior margin. The propodus is about 3 1/2
to 4 1/2 times the length of the dactylus. The carpus bears the usual movable
spine, while the merus as a rule shows 4 movable spines on the outer surface
near the posterior margin. The ischium also bears a spine. In the fifth leg
the dactylus is provided on the posterior margin with a row of many comb-
like arranged spinules. The propodus is almost 4 times as long as the dactylus.
The carpus bears one, the merus three movable spines.

The endopod of the lirst male pleopod is ovate, without any trace of
an appendix interna. In the second male pleopod the appendix masculina
is shorter than the appendix interna and bears an apical tuft of bristles.
The diaeresis of the uropodal exopod bears 8 to 11 spinules.

No ovigerous females are présent.

The présent species may be immediately distinguished from the related
forms by the shape of the rostrum, the large number of dorsal rostral teeth
of which 4 to 6 are placed behind the orbit, and by the shape of the telson.

The type lot was doubtfully identified by me in 1956 as C. gladii/era
J. Roux. Now that I hâve material of that species at my disposai it is clear
that the présent material forms an independent species.

Garidina lamiana n. sp. (fig. 14)

Madagascar Instituts :

— Ambodivoangy near Maroantsetra, Antongil Bay, NE Madagascar ; in torrent ;
1946 ; J. Millot. — 2 specimens.

Muséum Leiden :

— Forest between 20 and 30 km NW of Ambahoabé, Simianona River, N of Soanie-
rana, eastcoast of Madagascar; 8 December 1938; II. J. Lam & A. D. J. Mceuse. —
28 specimens (4 ovigerous).

The specimens are 9 to 17 mm long ; the ovigerous females ineasure
15 to 17 mm.

The rostrum is short and reaches about to the end of the basal segment
of the anlennular peduncle, in juvéniles it faits to attain the end of the eyes.
It is rather high, and is not armed, neither dorsally nor ventrally.

The antennal spine is rather broad and short, often showing as only
a mere angle. It stands below the lower orbital angle which is strongly
folded inward and therefore hidden from view. The pterygostomian angle
is rectangularly rounded and hardly produced forward.

The sixth abdominal somite is about 0.5 to 0.6 times as long as the cara¬
pace. The telson bears 4 or 5 pairs of dorsal spines. Its posterior margin

Source : MNHN, Paris

40

i. HOLTHUIS

is rounded. The outer pair of spinules is very short, measuring less than 1/4
of the length of the latéral spines, which are very long and strong. There
are 6 to 10 intermediate spines which are considerably shorter than the
laterals. The preanal carina is rounded, without spine.

The eyes are well developed with globular dark-pigmented cornea.

Fig. 14. — Caridina lamianu new species, from Simianona River, a, anterior nar»
body in latéral view ; b, tip of telson ; c, thiril maxilliped ; d, first pcreiopod • e second
pereiopod ; /, third pereiopod ; g, flfth pereiopod ; h, first pleopod of male’ • Y
part of diaeresis. a, c-g, x 16 ; b, i, x 32 ; h, x 50. ’ ’ outer

The antennular carina is dorsally produced to a high tooth-like élévation
The antennular peduncle is rather robust, with short segments. The stvlo-
cerite is sharp and pointed and fails to attain the end of the first segment
of the antennular peduncle. The antérolatéral looth of the first segment
is sharply pointed, it does not attain the middle of the second segment
The scaphocerite lias the final tooth rather broad, it is distinctly overreached
by the lamella. A distinct triangular tooth is placed at the external sid e
of the antennal peduncle near the base of the scaphocerite.

The. branchial formula is normal. Epipods are présent at the bases of
the lirst four pereiopod s. The third maxilliped reaches with 1/2 or 1/3 0 f
the distal segment beyond the final tooth of the scaphocerite, sometimes

Source : MNHN, Paris

E OF MADAGASCAR

•11

it barely attains that tooth. The distal segment is about as long as the
penultimate and bears some teeth in the distal half of the posterior margin.

The carpus of the first pereiopod is about twice as long as high and dis-
tinctly hollowed anteriorly. The carpus is as long as the merus and shorter
than the chela. The chela is very heavy, with the fingers about as long as
the palm. The second legs are slender. The carpus is 5 times as long as high,
it is longer than either merus or chela. The fingers of the chela are much
longer than the palm. The third pereiopod has 5 or 6 small denticles on the
posterior margin of the dactylus. The propodus is 3.5 to 4.5 times as long as
the dactylus. The carpus and ischium bear one, the merus three movable
spines near the posterior margin. The dactylus of the fifth leg is provided
with many comb-like arranged spinules on the posterior margin. The pro¬
podus is 3.0 to 3.5 times as long as the dactylus. The carpus bears one, the
merus two (sometimes three) and the ischium no movable spines near the
posterior margin.

The endopod of the first pleopod of the male is ovate and bears no
appendix interna. In the second male pleopod the appendix masculina is
about twice as long and twice as broad as the appendix interna, it bears
strong spines in the distal half and in two rows over the larger part of its
length. The diaeresis of the uropodal exopod bears 20 to 23 spinules.

The eggs are large, measuring 0.7 x 1.1-1.2 mm.

The species, in having no dorsal teeth on the rostrum, resembles C. i.
isa/oensis, but may be easily distinguished from that species by the high
number of spinules on the diaeresis.

This new species is named for Dr. H. J. Lam, Professor of Systematic
Botany at Leiden University and Director of the Rijksherbarium, who
together with Dr. Meeuse collected the type material. It is a pleasure for
me to be able in this way to express my high esteem for Dr. Lam and my
appréciation for his guidance in the field of taxonomy.

Caridina xiphias Bouvier (fig. 15, 16)

Caridina nilotica stylirostris Bouvier, 1925, p. 148, fig. 309.

Caridina nilotica xiphias Bouvier, 1925, p. 149, figs. 310-312 ; J. Roux,
1929, pp. 304-318.

Caridina gladiifera J. Roux, 1929, pp. 306, 319, fig. 16.

Madagascar Instjtute :

— Andranomaliza, East of Manakambahiny, eastern shore of Alaotra Lake, E Mada¬
gascar ; April 1956. — 2 specimens.

— Anove, Manambato, east coast of Madagascar N of Tamatave ; February 1955 ;
P. A. Robinson. — 1 specimen.

— Périnet, 25 km east of Moramanga, E Madagascar ; August 1946 ; J. Miliot. —
4 specimens (3 ovigerous).

— Périnet, in brook ; 3 Mardi 1950; J. Doucet. — 1 specimen.

— Périnet : 22 September 1954 ; E. Razafimandimby. — 2 specimens.

— Périnet ; March 1955. — 2 specimens.

— Périnet ; mountain torrent in forest, in pool at base of cascade ; 15 Decembcr 1955 ;
B. Stuckenberg. — 8 specimens (4 ovigerous).

— North of Anosibé, 30 km S of Moramanga ; in forest ; January 1951 ; R. Paulian.
— 4 specimens (1 ovigerous).

— Ifanadiana, torrent of Ranomafana, E Madagascar ; May 1951 ; R. Paulian. —
24 specimens.

Source : MNHN, Paris

42

.. B. HOLTHU1S

Austhian Madagascar Expédition :

— Source in Virgin forest near Mandraka, 68 km E of Tananarive ; 22 July 1958;
F.M. 86. — 12 specimens.

— Analamazaotra brook, Périnct; 28 July 1958; F.M. 102. — 1 specimen.

— Eflluent from Lac Vert, Périnct ; 23 July 1958; F.M. 105. — 3 specimens.

— Brook emptying in I.ac Vert, Périnet ; 28 July 1958; F.M. 109. — 18 specimens

— Brook emptying in Lac Rouge. Périnet : 23 July 1958: F.M. 111. — 10 specimens.

— Brook near Lac Rouge, Périnct ; 23 July 1958: F.M. 112. -— 6 specimens.

— Source in Virgin forest near Périnet; 28 July 1958; F.M. 113. — 1 specimen.

— Brook in Virgin forest, Périnct ; 31 July 1958;' F.M. 120. — 31 specimens (1 ovi-
gerous).

— Virgin forest near Périnet; 28 July 1958 ; T. 124. — 1 specimen.

Muséum Leiden :

— Beforona River, Périnet région. Moramanga district, Tamatave province, Central
Madagascar; 11 February 1961 ; native naine « orana »; Y. Therezien no. 10. — 1 ovige-
rous female.

— Brook at Périnet station, Moramanga district ; 12 May 1962 ; native name « patsa
Y. Therezien no. 12. — 13 specimens (2 ovigerous).

— Ambodiriana, Moramanga district ; 30 Octobér 1958 ; Y. Therezien no. 43, récolte
no. 12-14. — 1 specimen.

Analamazaotra River, Périnet, Moramanga district ; 29 July 1958; Y. Therezien
no. 44, récolte no. 6-11. — 9 specimens.

— Ampamaherana River. 1-ianarantsoa district, Fianaranlsoa province, F. Madagas¬
car ; 29 November 1961 ; native name «patsa •; Y. Therezien no. 15. — 13 specimens
(4 ovigerous).

The specimens are 7 to 26 mm long, the ovigerous females measure
21 to 26 mm.

The rostrum is variable in length. In adults it may overreach the scapho-
cerite by slightly less than half the length of that scale. In juvéniles it may
be much shorter and even fail to reach the base of the last segment of the
antennular peduncle. In juvéniles it usually is straight, in adults it may be
curved upwards in the distal part. The upper margin is unarmed distally,
even a subapical tooth is absent. The unarmed portion in adults may be un
to twice as long as the armed part ; in juvéniles the unarmed portion may be
as short as half the length of the armed part. The number of dorsal teeth
on the rostrum varies between 4 (in juvéniles) and 16 (in adults). 0 to 3 of
these teeth are placed behind the posterior limit of the orbit. The ventral
margin bears 0 (in very young specimens) to 12 teeth.

The lower orbital margin is distinctly separated from the antennal
spine winch is well developed and sliarply pointed. The pterygostomian
angle is rounded and somewhat forward produced.

The telson bears 3 to 5, usually 3 or 4, dorsal pairs of spines. The posterior
margin of the telson is rounded without a médian spine. The outer pair of
spines is very short, less than 1/4 of the length of the laterals. There are
5 to 9 intermediate spines, which are shorter, though not very much shorter
than the laterals. The preanal carina is blunt, without a spine.

The stylocerite reaches beyond the middle of the basal antennul ar
segment, but does not attain the end of it, though it often cornes quite close
The antérolatéral tooth of the basal segment is sharp and long ; in somé
cases it almost reaches the middle of the second segment, but usually fal] s
distinctly short of that. There is a distinct spine at the external side of the
antennal peduncle near the base of the scaphocerite.

The branchial formula is normal. Epipodites are présent at the bases
of the first four pereiopods.

The carpus of the first pereiopod is about as long as the merus, twic e

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

43

Fio. 15. — Caridina xiphias Bouvier, a-d, anterior part ol bodv in latéral view ; e-g, tip
ot telson ; h, i, outer part of diaeresis ; /, third maxillipcd ; k, I, endopod of flrst
pleopod of male, a, c, a, j, x 13 ; b, x 26 ; e, h, x 40 ; /, g, i, k, l, x 50. a, b, e, h,
specimen from brook near Lac Rouge, Périnet, F.M. 111 ; c, specimen from Périnet,
March 1955 ; d, I, g, i, i, I, specimen from Ifanadiana : k, specimen from Périnet,
F.M. 120.

or less than twice as long as high and shorter than the chela. The carpus
of the second pereiopod is longer than either merus or chela, it is 4.5 to
5 times as long as high. The dactylus of the third leg bears 5 or 6 spinules
on the posterior margin. The propodus is 4 to 4.5 times as long as the dactylus.
The propodus of the fifth leg is 3 to 4 times as long as the dactylus.

Source : MNHN, Paris

44

L. B. HOLTHIUS

The endopod of the fîrst pleopod of the male has no appendix interna
The diaeresis of the uropodal exopod bears 9 to 20 spinules (more in thé
adult tlian in the very young).

The eggs measure 0.5-0.7 x 0.9-1.2 mm.

The présent species superficially resembles Caridina nilolica very much
and for that reason has been treated by Bouvier (1925) as a subspecies of it
There can be not the least doubt, however, that the two forms constitutê

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

45

distinct species. The most important differential character in my opinion
being found in the shape of the endopod of tlie first pleopod of the male,
which in C. nilolica (as well as in C. longirostris) is provided with a well
developed appendix interna, while such an appendix is entirely absent in
C. xiphias. C. xiphias furthermore is readily distinguishable from C. nilolica
by the lack of subapical teeth on the rostrum (in C. nilotica these teeth
may be absent in abnormal or damaged specimens), the more produced
pterygostomian angle of the carapace and the fewer teeth on the rostrum.

As shown by my material several of the characters of this species show
a great variation, which is either individual or due to âge. A comparison
of the characters with the original descriptions of both Caridina nilotica
xiphias Bouvier, 1925, and C. n. stylirostris Bouvier, 1925, shows that these
forms both belong to the présent species. Bouvier’s specimens of C. n.
xiphias were immature, as stated by that author, those of C. n. stylirostris
were adult. The names xiphias is retained here for the species.

J. Roux (1929) dealt with C. n. xiphias and described the new speciess
Caridina gladii/era. As Roux himself already pointed out his new specie
has many characters in common with C. n. xiphias. The most conspicuous
différence being the greater number of spines on the diaeresis of the uropodal
exopod. As shown by the présent material this différence is of little value
and Caridina gladii/era in my opinion must be considered a synonym of
C. xiphias as also the otlier characters do not fall out of the range of variation
shown by the species.

The character of the piesence or absence of subapical dorsal teeth on
the rostrum, though very useful for the distinction of C. xiphias and C. nilo¬
lica, cannot be always relied upon. In some specimens of C. nilolica these
spines are absent probably due to some accident, while in C. xiphias 1 saw
once a specimen with an indication of a subapical spine. The type specimens
of Caridina voeltzkowi Lenz (1910, p. 569) were described by Lenz as having
no subapical dorsal teeth, still the few other characters of the specimens
provided by Lenz (e.g., the rostral formulae) show that the specimens are
more probably C. nilotica ; this is confirmed by J. Roux (1929, p. 302)
who evidently examined Lenz’s specimens and identified them as C. nilotica
paucipara.

Caridina xiphias is only known from Madagascar. The type locality is
Didy, Ambatondrazaka district, province Moramanga, E Madagascar.
The localities whence the species has been reported in the literature are :
Madagascar (Bouvier, 1925), Didy (Bouvier, 1925 ; J. Roux, 1929),
Périnel, 25 km east of Moramanga, E Madagascar (J. Roux, 1929), Namoroka
and Kapiloza Rivers, Ambongo, W Madagascar (J. Roux, 1929). It is
possible that J. Roux’s specimens from W Madagascar do not belong here.
Juvénile specimens of this species and those of C. isaloensis grandidieri
often resemble each other so strongly that it is impossible to tell them apart.

Caridina hova Nobili (fig. 17)

Caridina hova Nobili, 1905, p. 1, 3 figs. ; Bouvier, 1925, p. 186, fig. 390;

Decary, 1950, p. 167.

Austrian Madagascar Expédition :

— Mandromodromotra brook, 20 km N of Fort-Dauphin, SW Madagascar; 11 Sep-
tembcr 1958 ; F.M. 217. — 10 specimens (2 ovigerous).

Source : MNHN, Paris

46

L. B. HOLTHUIS

_Source in marsh20kmNof Fort-Dauphin; 11 Septemberl958; F.M.220.— 1 specimen.

Muséum Leiden :

— Mandromodromotra brook, Piste Elandy, Fort-Dauphin district, Tuléar province
SE Madagascar ; 11 September 1958 ; Y. Y. Therezien no. 36, récolte no. 46. — 2 spécimens.

The specimens are 8 to 18 mm long. The ovigerous females measure
17 and 18 mm.

The rostrum reaches to the base or the end of the second segment of
the antennular peduncle. It is straight and bears dorsally 14 to 20 teeth,
3 of which are placed behind the posterior limit of the orbit. These teeth

are striking by their large size and form the most conspicuous characteristic
of the species. The teeth are evenly distributed over the rostrum. The lower
margin bears 3 or 4 distinct teeth. The lower orbital angle is distinct and
acutely rounded. The antennal spine is strong and placed some distance
below the orbital angle. The pterygostomian angle is rectangularly rounded.

The telson bears 4 or 5 pairs of dorsal spines. The posterior margin is
rounded. The outer spines are less than 1/4 as long as the laterals. The inter-
mediate spines are 5 to 7 in number and are about half as long as the laterals.
The pre-anal carina is bluntly rounded without a spine.

The eyes are well developed. The stylocerite reaches beyond the middh»
of the basal segment of the antennular peduncle but does not attain the

Source : MNHN, Paris

THE ATYIDAE OF MADAGASCAR

47

end of the segment. The antérolatéral tooth of the segment is sharp, but
does not reach the middle of the second segment. The antennal peduncle
bears a strong spine in its external part near the base of the scaphocerite.

The branchial formula is normal. Epipodites are présent at the bases
of ali 4 pereiopods. The distal segment of the third maxilliped is about
as long as the penultimate. The carpus of the first pereiopod is fully 2.5 times
as long as high, it is about as long as the merus and only slightly shorter
than the chela. The second leg has the carpus 6 to 8 times as long as high,
being longer than either merus or chela ; the fingers of the chela are slender
and distinctly longer than the palm. The dactylus of the third pereiopod
bears about 6 teeth on the posterior margin ; the propodus is about 5 times
as long as the dactylus. The propodus of the fifth leg is 4 times as long as
the dactylus, which bears numerous comb-like arranged spinules on the
posterior margin.

The endopod of the first pleopod of the male bears no appendix interna.
The diaeresis of the uropodal exopod is provided with 15 to 17 spinules.

The eggs measure 0.65-0.7 x 1.1 mm.

Caridina hova is so far only known from SE Madagascar. Nobili (1905)
reported it from Fort-Dauphin and since that time no new finds of the
species hâve been published. The présent material also originates from the
Fort-Dauphin area.

Caridina brevirostris brevipes Bouvier
Caridina brevirostris brevipes Bouvier, 1925, p. 230.

The status of this form, of which no material has been examined by
me is dubious. Bouvier only provided a very short description of it, based
on two specimens from the Bay of Amposindova, opposite Nosy Bé, NW
Madagascar.

Genus ATYA Leach, 1816

So far only one species of this genus is known from Madagascar.

Atya pilipes Newport

Atya pilipes Newport, 1847, p. 160.

Atya brevirostris Coutière, 1900, p. 1267.

Atya serrala Bouvier, 1925, p. 294.

Muséum Leiden :

— Rivulct of Besokatra, Diégo-Suarez district, Diégo-Suarez province, N Madagascar;
25 April 1959; Y. Therezien no. 40, récolte no. 103. — 1 female.

The specimen is 40 mm long. It agréés quile well with the descriptions
given in the literature of this species. It belongs to the form which Bouvier
(1925, p. 294) described under the name Atya serrata Bâte.

Previous records of the species from Madagascar are the following :
Cap d’Ambre, northpoint of Madagascar (Coutière, 1900), and the island
Sainte-Marie, E Madagascar (Bouvier, 1925).

Atya pilipes has a wide distribution throughout the Indo-West Pacific
région, viz., from Madagascar and the Seychelles to Polynesia.

Source : MNHN, Paris

48

!.. B. HOLTHUIS

REFERENCES

Bouvier (E. L.), 1904. — Crevettes de la famille des Atyidés : espèces qui font partie
des collections du Muséum d’histoire naturelle. Bull. Mus. Hisl. nat. Paris vol in
pp. 129-138. ’

— 1905. — Observations nouvelles sur les Crevettes de la famille des Atvidés R
sa. France Belg., vol. 39, pp. 57-134, fig. 1-26.

—- 1912. — Les Caridines de l’île Maurice, d’après les envois faits au Muséum par M lo
Dr 1..-G. Barbeau. Bull. Mus. Hist. nat. Paris, vol. 18, pp. 291-300.

— 1912 o. — Sur la classification du genre Caridina et les variations extraordinaires
d’une espèce de ce genre, la Caridina brevirostris Stimpson. C. B. Acad. Soi. Pari*

vol. 154, pp. 915-922.

— 1913. — Les Caridines des Seychelles. The Percy Sladen Trust Expédition to the

Indian Océan in 1905, under the leadership of Mr. J. Stanley Gardiner. Trans I inn
Soc. Lond. Zool., ser. 2, vol. 15, pp. 447-472, pl. 27-29. ' *

— 1919. -— Quelques espèces nouvelles de Caridines. Bull. Mus. Hist. nat. Paris vol •>=.

pp. 330-335. ’

- 1925. — Recherches sur la morphologie, les variations, la distribution géosranhim.a
des Crevettes de la famille des Atyidés. Encycl. en/., ser. A, vol. 4, pp. 1-370, fig.1-7Ï6

Coutière (H.), 1899. — Sur quelques Macroures des eaux douces de Madagascar (Vovuro
de M. G. Grandidier). Bull. Mus. Hisl. nal. Paris, vol. 5. un. 382. 383. ' Ke

— 1900. — Sur quelques Macroures des e;

Paris, vol. 130, pp. 1266-1268.

Decahy (R.), 1950. — La faune malgache, s
gènes, pp. 1-236, fig. 1-22.

Edmondson (C. H), 1935. — New and rare Polynesian Crustacea. Occ. Pap. BishoD

Honolulu, vol. 10 pt. 24, pp. 1-38, textflgs. 1-11, pl. 1, 2. •

Face (L.), 1946. — Sur une Caridine nouvelle cavernicole de Madagascar, Caridina miern
phthalma, nov. sp. Bull. Mus. Hisl. nal. Paris, ser. 2, vol. 18, pp. 324-327, flg. i

Gordon (I.), 1930. — African Freshwater Prawns of the Spccies Caridina nilotica (Rou'vl'
'vith^Spécial Référencé to the Nile Basin. Proc. zool. Soc. Lond., 1930, pp. 33-5o’

Holthuis (L. B.), 1956. — An Enumération of the Crustacea Decapoda Natantin inh«
biting suhtcrrancan Waters. Vie et Milieu, vol. 7, pp. 43-70. na ~

— 1956 a. — The troglobic Atyidae of Madagascar (Crustacea Decapoda Natanti-,\
Mém. Inst. sci. Madagascar, ser. A, vol. 11, pp. 97-110, flg. 1-4.

Kubo (L), 1938. — On the .lapanese Atyid Shrimps. Journ. Imp. Fish. Inst TnL-,.„
vol. 33, pp. 67-100, flg. 1-24. ' Aj/ °>

s, vol. 5, pp. 382, 383.
x douces de Madagascar. C. R. Acad. Sci.

n rôle dans les croyances et les usages indi-

! (H.), 1905.

Herrn Prof. Dr. ,

der Reisen in Madagaskar und Ostafrika i. .

Senckenb. naturj. Ges., vol. 27, pp. 341-392, pl. 47, 48.

iom . von Madagaskar, Ostafrika und Ceylon. In : Voeltzkow ( A \

i den .iahren 1903-1905 mit Mitteln der Hermann und |;ii„

. -.9. nn V10_r,7fi c-USe

- Ostafrikanische Dekapoden und Stomatopoden gesammelt von
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- 1910. — Crustaceen v

Reise in Ostafrika in _ _ _

geb. Ileckmann Wentzei-Stiftung ausgeführt, vol. 2, pp. 539-576.

Man (J. G. de), 1892. -— Decapodcn des Indischen Archipels. In : Weber (M.), Zoologisch
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vol. 14, pp. 225-264, pl. 7-10. Us -

Milne Edwards (H.), 1837. — Histoire Naturelle des Crustacés, comprenant l’anatomi
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Newport (G.), 1847. — Note on the genus Atya of Leach, with descriptions of fm„-
apparently new Species, in the Cabinets of the British Muséum. Ann Man „ ,
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Nobili (G.), 1905. Descrizione di una nuova Caridina del Madagascar. Boll Vf..

Zool. Anal. comp. Torino, vol. 20 n. 499, pp. 1-3, 3 flg. ’ Us -

Rathbun (M. J.), 1906. — The Brachvura and Macrura of the Hawaiian Islands R„n
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Roux (J.), 1929. — Crustacea. III. Atyidae. In : Petit (G.). Contribution à l’étude de la
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— 1934. - Macroures d’eau douce de Madagascar et des îles voisines (Palémoni.iz

et Atyidés). Faune Colon. Franç., vol. 5, pp. 529-547, flg. 1,2. aés

Roux (P.), 1833. — Lettre relative à divers
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s Coquilles, Crustacés, Insectes, Reptiles «♦
nal., vol. 28, pp. 72-78, pl. 7. et

Pierre André imp., 244 Boulevard Raspai:

Dépôt légal : 3 e Trimestre 1964

Achevé d’imprimer le 20 Juillet 1965

Source : MNHN, Paris