The freshwater atyid shrimps of the genus Caridina (Crustacea Decapoda Caridea) from Lake Lindu, Central Sulawesi, Indonesia

THE FRESHWATER ATYID SHRIMPS OF THE GENUS
CARIDINA (CRUSTACEA: DECAPODA: CARIDEA)
FROM LAKE LINDU, CENTRAL SULAWESI, INDONESIA

ANNAWATY

SEKOLAH PASCASARJANA
INSTITUT PERTANIAN BOGOR
BOGOR
2014

PERNYATAAN MENGENAI DISERTASI DAN
SUMBER INFORMASI SERTA PELIMPAHAN HAK CIPTA*
Dengan ini saya menyatakan bahwa disertasi berjudul The freshwater atyid
shrimps of the Genus Caridina (Crustacea: Decapoda: Caridea) from Lake Lindu,
Central Sulawesi, Indonesia adalah benar karya saya dengan arahan dari komisi
pembimbing dan belum diajukan dalam bentuk apa pun kepada perguruan tinggi
mana pun. Sumber informasi yang berasal atau dikutip dari karya yang diterbitkan
maupun tidak diterbitkan dari penulis lain telah disebutkan dalam teks dan
dicantumkan dalam Daftar Pustaka di bagian akhir disertasi ini.
Dengan ini saya melimpahkan hak cipta dari karya tulis saya kepada Institut

Pertanian Bogor.
Bogor, Agustus 2014
Annawaty
NIM G362090011

RINGKASAN
ANNAWATY. Udang air tawar Genus Caridina (Crustacea: Decapoda: Caridea)
dari Danau Lindu, Sulawesi Tengah, Indonesia. Dibimbing oleh BAMBANG
SURYOBROTO, ACHMAD FARAJALLAH, DAISY WOWOR, dan DEDI
SETIADI.
Udang air tawar endemik Danau Lindu, Caridina linduensis pertama kali
dideskripsi lebih dari satu abad yang lalu oleh J. Roux berdasarkan koleksi milik
naturalis dari Swiss, Paul dan Fritz Sarasin (Roux 1904). Dalam tulisan ini
dideskripsi ulang C. linduensis berdasarkan specimen baru yang diperoleh dari
Danau Lindu yang merupakan type locality dari spesies tersebut. Hasil penelitian
ini juga mengungkapkan adanya dua spesies udang air tawar endemik lain yang
belum pernah dideskripsi sebelumnya, yaitu Caridina sp A dan Caridina sp B,
kedua spesies tersebut dideskripsi dan dibuat ilustrasinya dalam tulisan ini.
Ketiga spesies Caridina danau Lindu dapat dibedakan terutama berdasarkan
bentuk rostrum dan struktur mandibulanya. Caridina linduensis memiliki rostrum

yang panjang dengan ujung rostrum mencapai atau sedikit melewati ujung segmen
ketiga antennular pedunclenya. Spesies ini memiliki rostrum yang paling panjang
dibandingkan dengan kedua spesies Caridina yang lain. Rostrum Caridina sp B
lebih pendek dengan ujung rostrum melewati ujung segmen basal tetapi tidak
pernah mencapai ujung segmen kedua dari antennular peduncle. Caridina sp A
dapat dibedakan dari kedua spesies lainnya berdasarkan bentuk rostrum yang
sangat pendek, hanya mencapai ujung distal segmen basal dari antennular
peduncle. Bentuk rostrum yang sangat pendek ini membuatnya sangat berbeda
dari kedua caridina yang lain. Disamping itu Caridina sp A memiliki ukuran
tubuh yang lebih besar dibanding kedua spesies yang lain. Panjang carapas
individu betina bertelur Caridina sp A mencapai 6.8 mm sementara C. linduensis
hanya 4.9 mm dan 5.4 mm pada Caridina sp B. Struktur mandibula pada Caridina
sp A dan Caridina sp B memiliki kemiripan dengan adanya 6 gigi pada incisor
process dengan 2 deret granula yang tidak beraturan di antara gigi, molar process
lurus. Struktur ini sangat berbeda dengan C. linduensis, yang memiliki lebih
banyak gigi pada incisor processnya yaitu 16-18 gigi dengan struktur molar
process yang melengkung.
Ketiga spesies Caridina tidak hanya berbeda secara morfologi, tetapi juga
berbeda dalam pemilihan habitat serta distribusinya. C. linduensis ditemukan
menggantung di akar tanaman air di daerah littoral danau dan ditemukan juga di

permukaan serasah dan ranting ranting kayu yang sudah lapuk di tepi outlet danau
yang memiliki arus lambat cenderung stagnan. Caridina sp B ditemukan di bagian
danau yang berada tidak jauh dari inlet, dan juga ditemukan di inlet dengan air
keruh dengan arus yang lambat cenderung stagnan. Spesies ini menggantung di
akar tumbuhan pakis atau tanaman air yang terdapat di tepi danau atau sungai..
Caridina sp A hanya dapat ditemukan di sungai ataupun selokan kecil berair
jernih dengan substrate baru kerikil dan kerakal serta aliran air yang cukup deras.
Spesies ini melimpah di sungai-sungai yang terletak di bagian barat danau,
dimana kemiringan lahan berkisar antara 15 hingga 40 derajat.
Distribusi Caridina di Danau Lindu dan sungai-sungai sekitarnya
menunjukkan tidak adanya tumpang tindih di antara satu spesies dengan spesies

yang lain, dan tampaknya distribusinya dipengaruhi oleh temperatur dan adanya
ikan-ikan introduksi seperti mas (Cyprinus carpio) dan mujair (Oreochromis
mossambicus), serta adanya udang introduksi (Macrobrachium lanchesteri).
Fauna akuatik introduksi ini, baik yang disengaja maupun tidak, merupakan
ancaman serius bagi kelangsungan hidup Caridina terutama spesies yang hidup di
danau karena berpotensi sebagai predator maupun kompetitor bagi Caridina.
Semua spesies Caridina Danau Lindu memiliki jumlah telur yang relatif
sedikit (26–47 telur) dengan ukuran telur yang cukup besar (diameter lebih dari 1

mm). Hal ini mengindikasikan sifat mereka sebagai spesies land-locked yang
berarti tidak lagi membutuhkan air payau/asin untuk menyelesaikan siklus
hidupnya karena mereka sepenuhnya menghabiskan seluruh hidupnya di air tawar.
Perpaduan antara tipe land-lock dengan barrier ekologi dan fisik yang
membatasi penyebaran Caridina nampaknya juga berkontribusi terhadap
berkurangnya aliran gen di antara satu populasi dengan populasi yang lain. Hal ini
dapat menjelaskan tingginya derajat endemisitas Caridina di Danau Lindu. Di
samping itu dibandingkan dengan Caridina yang terdapat di Danau Poso dan
Kompleks Danau Malili, tidak ditemukan adanya tumpang tindih spesies antara
satu danau dengan danau yang lain. Semua Caridina yang terdapat di masingmasing sistem danau adalah endemik terhadap masing-masing sistem danau.
Kata kunci: Caridina, spesies baru, distribusi, Danau Lindu, Sulawesi

SUMMARY
ANNAWATY. The freshwater atyid shrimps of the genus Caridina (Crustacea:
Decapoda: Caridea) from Lake Lindu, Central Sulawesi, Indonesia. Supervised by
BAMBANG SURYOBROTO, ACHMAD FARAJALLAH, DAISY WOWOR,
and DEDI SETIADI.
The freshwater atyid shrimps from Lake Lindu, Caridina linduensis are
described for the first time by J. Roux in 1904. He described the species based on
the specimen collection of Paul and Fritz Sarasin. After neglected for more than a

century, here, we redescribe and figure this poorly known species based on new
material from its type locality, Lake Lindu. Two new species, Caridina sp B and
Caridina sp A are also found in this lake and they are described and illustrated.
The three endemic species have different morphological characters mainly
in rostrum shape and mandible structure. Rostrum on C. linduensis relatively long
with the rostrum tip reaching near to or slightly beyond end of the third segment
of antennular peduncle. This species have the longest rostrum compared with the
other species. Rostrum of Caridina sp B is rather short which is tip of rostrum
overreach the end of basal segment, but never reach end of second segment of
antennular peduncle, while Caridina sp A have a very short rostrum with
maximum reach of the end of basal segment of antennular peduncle. Caridina sp
A have a shorter rostrum among the Lindu’s species. The shortest rostrum in
Caridina sp A make this species very different from the two Caridina species
found in the Lake Lindu and its catchment area. Caridina sp A also have larger
body size of ovigerous females with maximum carapace length 6.8 mm while only
4.9 mm in C. linduensis and 5.4 mm in Caridina sp B. The second morphological
difference of three Caridina species is its mandible structure. The mandibles of
Caridina sp B and Caridina sp A are similar, i.e. the incisor process has 6
irregular teeth with 2 very fine granulated rows between the teeth, and the molar
process is straight. On the other hand, the mandible of C. linduensis is quite

different; the incisor process has 16–18 irregular teeth, and the molar process is
truncated.
The Caridina from Lake Lindu not only different in their morphology but
also in their habitat preferences. C. linduensis is a true lake inhabitant, Caridina
sp B can be found both in the lake itself and associated streams while Caridina sp
A is an obligate stream species. There is no overlapping distribution among the
species.
Caridina sp A is abundant in streams and ditches with moderate flow
running water gravel–cobble substrate. It is mainly spread in the western part of
the lake where slope varies between 15 and 40%. Both Caridina sp B and C.
linduensis can be found in lake and streams with very slow current to almost
stagnant water, muddy sand substrate and associated with roots of water plants
and leaf litter. However, Caridina sp B is never occurred together with C.
linduensis and they are less abundant compare to Caridina sp A. Distribution of
Caridina spp in Lake Lindu is probably affected by the temperature of their
habitats and the occurrence of introduced fish such as Mozambique tilapia
(Oreochromis mossambicus), common carp (Cyprinus carpio), and an alien
riceland prawn (Macrobrachium lanchesteri). It is the first record for M.

lanchesteri present in Lake Lindu.These exotic fish and the alien shrimp can have

the potency become predators and or competitors for the Caridina spp.
The less egg number (26–47) and the large egg size (more than 1 mm in
diameter) of the all Caridina species from Lake Lindu system indicated they are
probably land-locked species which are complete their entire life cycle in
freshwater. In combination with their land-locked species type and ecological and
physical barrier, the Lindu’s Caridina distribution is restricted within the lake
system. The environmental condition prevents Caridina to disperse into other
area. It is also associated with the reduction of dispersal ability and gene-flow
among the populations, which may explain the high degree of endemism in
Caridina of Lake Lindu. There is no overlap of Caridina species among Lake
Lindu, Lake Poso and Malili lake system. Most of the species which belong to the
lake system are endemic to lake system itself.
Keywords: Caridina, new species, distribution, Lake Lindu, Sulawesi

© Hak Cipta Milik IPB, Tahun 2014
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THE FRESHWATER ATYID SHRIMPS OF THE GENUS
CARIDINA (CRUSTACEA: DECAPODA: CARIDEA)
FROM LAKE LINDU, CENTRAL SULAWESI, INDONESIA

ANNAWATY

Disertasi
sebagai salah satu syarat untuk memperoleh gelar Doktor
pada Program Studi Biosains Hewan

SEKOLAH PASCASARJANA
INSTITUT PERTANIAN BOGOR
BOGOR
2014

Penguji pada Ujian Tertutup:

1. Prof Dr Mulyadi, MSc
Professor riset Pusat Penelitian Biologi,
Lembaga Ilmu Pengetahuan Indonesia
2. Prof Dr Ir MF Rahardjo, DEA
Guru Besar Fakultas Perikanan dan Kelautan IPB

Penguji pada Ujian Terbuka:
1. Prof Dr Ramadhanil Pitopang, MSi
Guru Besar Fakultas MIPA Universitas Tadulako
2. Prof Dr Ir MF Rahardjo, DEA
Guru Besar Fakultas Perikanan dan Kelautan IPB

Judul Disertasi

Nama
NIM

: The freshwater atyid shrimps of the genus Caridina
(Crustacea: Decapoda: Caridea) from Lake Lindu, Central
Sulawesi, Indonesia

: Annawaty
: G362090011

Disetujui oleh
Komisi Pembimbing

Dr Bambang Suryobroto
Ketua

Dr Ir Achmad Farajallah, MSi
Anggota

Dr Ir Daisy Wowor, MSc
Anggota

Prof Dr Ir Dede Setiadi, MS
Anggota

Diketahui oleh

Ketua Program Studi
Biosains Hewan

Dekan Sekolah Pascasarjana

Dr Ir Rd Roro Dyah Perwitasari, MSc

Dr Ir Dahrul Syah, MScAgr

Tanggal Ujian: 11 Agustus 2014

Tanggal Lulus:

PRAKATA
Puji dan syukur penulis panjatkan kepada Allah SWT atas segala karuniaNya sehingga karya ilmiah ini berhasil diselesaikan. Tulisan yang berjudul “The
freshwater atyid shrimps of the genus Caridina (Crustacea: Decapoda: Caridea)
from Lake Lindu, Central Sulawesi, Indonesia” ini mengungkapkan adanya
penemuan dua spesies baru udang air tawar dari danau Lindu yaitu Caridina sp A
dan Caridina sp B. Tulisan ini juga mendeskripsikan kembali Caridina linduensis,
udang air tawar endemik Danau Lindu yang merupakan satu-satunya udang air
tawar yang pernah dilaporkan dari Danau Lindu lebih dari 100 tahun yang lalu.
Tulisan ini mengungkapkan juga adanya pereferensi habitat dan distribusi yang
berbeda pada ketiga spesies Caridina tersebut.
Terima kasih penulis ucapkan kepada Dr Bambang Suryobroto, Dr Ir
Achmad Farajallah, MSi, Dr Ir Daisy Wowor, MSc dan Prof Dr Ir Dede Setiadi,
MS selaku komisi pembimbing. Ucapan terima kasih juga saya sampaikan kepada
seluruh staf pengajar di Program Studi Biosains Hewan serta teknisi di
Laboratorium Molekuler Program Studi BSH. Kepada tenaga lokal di Danau
Lindu serta mahasiswa Jurusan Biologi Universitas Tadulako yang telah
membantu selama proses sampling di Danau Lindu, penulis juga mengucapkan
terima kasih. Kepada rekan-rekan seprofesi di Jurusan Biologi Universitas
Tadulako serta rekan-rekan di Himpunan Mahasiswa Pascasarjana Sulawesi
Tengah, saya juga ucapkan terima kasih atas dukungannya.
Terima kasih penulis sampaikan kepada Dekan Fakultas MIPA Universitas
Tadulako dan Rektor Universitas Tadulako yang telah memberi izin untuk
melanjutkan pendidikan S3, Dirjen Dikti Kementerian Pendidikan dan
Kebudayaan atas beasiswa BPPS dan beasiswa Sandwich-like. Penulis juga
mengucapkan terima kasih kepada Dr Kristina von Rintelen yang telah
mengirimkan deskripsi asli Caridina linduensis dan Prof Rudolf Meier yang telah
membimbing penulis selama mengikuti program Sandwich-like di Evolutionary
Biology Laboratory, National University of Singapore.
Kepada rekan seangkatan 2009 Dr. Irma Shita Arlyza dan Puji Rianti MSi.
serta seluruh angkatan 2011 S3 BSH: Andy Darmawan, MSi, RR. Sri Catur
Setyawaningsih MSi, FX Widadi Padmarsari Soetignya MSi dan Yuliadi Zamroni
M.Si. penulis juga mengucapkan terima kasih atas diskusi dan dukungannya
selama penulis menyelesaikan penulisan disertasi.
Ungkapan terima kasih juga disampaikan kepada kedua orangtua penulis,
bapak dan ibu mertua, seluruh keluarga besar penulis, serta suami penulis, Zulfan
Hasibuan MPd atas segala dukungan dan kesabarannya selama penulis mengikuti
pendidikan doktor.

Bogor, Agustus 2014
Annawaty

DAFTAR ISI
DAFTAR TABEL

xi

DAFTAR GAMBAR

xi

1 INTRODUCTION

1

2 LITERATURE STUDY

2

3 MATERIAL AND METHODS
Study area
Collecting specimen
Environmental Data Analysis
Morphometric Measurements for Taxonomical Analysis

3
3
4
4
5

4 RESULTS AND DISCUSSION
Results
Discussion

6
6
25

5 CONCLUSION

29

REFERENCES

29

RIWAYAT HIDUP

33

DAFTAR TABEL

Table 1 Distribution of Caridina species collected in Lake Lindu system …..21
Table 2 Distribution of Caridina species in Lake Lindu, Lake Poso and Malili
lake system ……...…………………………………………………..24

DAFTAR GAMBAR
Figure 1 Morphology of Caridea shrimp ............................................................ 2
Figure 2 Location of the Lake Lindu system in Sulawesi Island (right)
and overview of the sampling sites in the lake (left). Circle refer to
the sampling site ........................................................................................... 4
Figure 3 Line drawing of Caridina linduensis Roux, 1904. ............................... 7
Figure 4 Line drawing of Caridina linduensis Roux, 1904. ............................... 8
Figure 5 Line drawing of Caridina sp B........................................................... 13
Figure 6 Line drawing of Caridina sp B........................................................... 14
Figure 7 Line drawing of Caridina sp A. ......................................................... 18
Figure 8 Line drawing of Caridina sp A. ......................................................... 19
Figure 9 Map of the Lake Lindu system, showing distribution of a)
Caridina linduensis b) Caridina sp A and c) Caridina sp B, presence
and absence as indicated. ............................................................................ 23
Figure 10 Habitat of Caridina in Lake Lindu system ....................................... 24

1 INTRODUCTION
Freshwater atyid shrimps are widely distributed in Indonesia. They can be
found in various water bodies such as lake, pond, river, stream and ditch, both in
surface and underground waters (Wowor et al. 2004). The shrimp habit as
scavengers or detritus feeders make this family very important in ecosystem to
recycle organic material. Leaf decay rate in the tropical stream was significantly
greater in the presence of shrimps than in their absence (March et al. 2001).
The shrimps are characterized by having chelipeds with setae on the tip of
the fingers which are used for filtering small aquatic organisms or scraping
detritus during feeding (Fryer 1977). The atyid shrimp is consisted of 469 species
dominated by the genus Caridina. The genus has 290 species members (De Grave
and Fransen 2011). In Indonesia, there are 62 species of atyid shrimps and 52 are
reported to be found in Sulawesi and nearby islands (De Grave and Fransen, 2011;
Klotz and von Rintelen 2013). The majority of the species are belong to the genus
Caridina with more than half of them are lacustrine and endemic to the island.
In the last ten years, the study of the lacustrine Caridina spp in Sulawesi has
been focused in taxonomy and evolution of the species. The studied lakes are
Poso and Malili lakes system which were considered by Schön and Martens
(2004) as ancient lakes. Recently, the ancient status of the lakes was confirmed
both in terms of their estimated age as well as their fauna (von Rintelen et al.
2012). It is agree with the definition of ancient lake which is usually perceived as
more or less long‐lived lake that often associated with both high species numbers
and endemicity in various systematic groups (Albrecht 2012).
Before 2006, there are only nine species known from the Lake Poso and
Malili lakes system. Recent studies record 12 new Caridina species from the lakes,
viz. C. caerulea, C. dennerli, C. glaubrechti, C. holthuisi, C. longidigita, C.
mahalona, C. parvula, C. profundicola, C. schenkeli, C. spongicola, C. striata,
and C. woltereckae (Zitzler and Cai 2006; Cai and Wowor 2007; Cai et al. 2009;
Rintelen and Cai 2009).
Although atyid shrimps from Lake Poso and the Malili Lake system has
been getting considerable attention, the fauna of Lake Lindu, has been more
neglected, with only one endemic species known thus far, Caridina linduensis
Roux, 1904. The specimens of this species were collected for the first time by
Swiss naturalists Paul Benedict Sarasin and Karl Friedrich Sarasin during the
1901–1903 Sulawesi Expedition.
During year 2011 field work, Annawaty and Wowor (2012) came across
two other undescribed Caridina species from the Lake Lindu, i.e. Caridina sp A
and Caridina sp B. The three Caridina species can be distinguished by the shape
of the rostrum and the structure of mandible (Annawaty and Wowor, submitted
manuscript).
In this study, C. linduensis is re-described, and the two new species are
described. Based on their observation, the Lindu’s Caridina not only differ in
their morphology but also in their habitat preferences. Therefore, to reveal the
habitat preferences and to map the distribution of the Caridina spp in Lake Lindu
and its catchment area become the other objective of this study.

2

2 LITERATURE STUDY
Freshwater shrimp belong to the order Decapoda, infraorder Caridea which
is characterized by the second abdominal pleuron (lateral plate) greatly expanded,
pear-shaped and overlapping posterior part of first pleuron and anterior part of
third pleuron (Chan 1998). Caridea shrimp consists of approximately 3438 species
in 35 families, one of the most speciose freshwater family is the Atyidae which is
consisted of 469 species and more than half of them belongs to the genus
Caridina (de Grave and Fransen 2011). The Family Atyidae are characterised by
unique feeding chelipeds, with the complex brushes on the first and second
pereiopods (Fig. 1) filtering out suspended matter or sweeping up microbial films
(Fryer 1977).
The study of the freshwater atyid shrimp fauna of Sulawesi was began more
than a century ago when the first atyid species, Atya wyckii described from the
Lake Tondano, North Sulawesi. The species was described by Hickson on 1888
(Klotz et al. 2007).
According to de Grave et al. (2008) half of 28 species of atyid shrimp
reported from Sulawesi are endemic to the island. The number of species
increased rapidly in the last ten years mainly for the lacustrine species. Cai and
Wowor (2007) described C. longidigita and re-described 3 other species i.e. C.
sarasinorum, C. ensifera and C. acutirostris from Lake Poso. The next 2 year, von
Rintelen and Cai (2009) described two new species from the lake, C. caerulea and
C. schenkeli and increase the number of Caridina species in the lake to become 6.
The other lake which is received considerable attention is Malili lakes
system. The lake system composed of three major lakes that directly connected,
Matano, Mahalona, and Towuti and two smaller satellite lakes, Lake Lontoa (or
Wawantoa) and Lake Masapi. There are several species described from 2006 i.e.
C. spongicola (Zitzler and Cai 2006), C. woltereckae and C. mahalona (Cai et al.
2009), C. dennerli, C. glaubrechti, C. holthuisi, C. parvula, C. profundicola and C.

Figure 1 Morphology of Caridina shrimp (modified from von
Rintelen and Cai 2009)

3
striata (von Rintelen and Cai 2009).
Caridina linduensis is the only atyid shrimp reported from Lake Lindu so
far. The description of the species based on the collection of Swiss naturalis, Paul
and Fritz Sarasin during the 1901–1903 Sulawesi Expedition (Roux 1904). Lake
Lindu located in the central highlands of Sulawesi (Lukman 2007) at an altitude of
982 m asl. This tectonic lake is found along a strike-slip fault zone, the Palu-Koro
Fault, formed from convergence of three major tectonic plates, i.e. Pacific, IndoAustralia and Eurasia. This fault zone is a fast slipping area but with a relatively
low level of seismicity (Bellier et al. 2001).
The age of Lake Lindu was estimated by Sarasin and Sarasin (1905) based
on the lake mollusk fauna, and it is believed that geologically, it is a very recent
structure that was formed through the sinking of a part of a mountain range during
the Pleistocene. However, the exact age of Lake Lindu is still not known. Lake
Lindu is drained by the Rawa River towards the north and in a point turns left to
join the Sopu River through a long, deep gorge. The confluence of Rawa and the
Sopu rivers marks the start of the Gumbasa River (Deschamps and Turland 2001)
which runs into Palu River and empties to Palu Bay.

3 MATERIAL AND METHODS
Study area
The Lake Lindu system (01°16’–01° 23’ S, 120°1’–120°11’E) is located in
the Lindu enclave in Lore Lindu National Park, Sulawesi (previously Celebes),
Indonesia. This relatively small lake covered an area of 34.5 km2 with 72.6 m
maximum depth, drain to the north via Rawa River, the only outlet for Lake Lindu,
to Palu Bay (Lukman 2007). The lake is fed by several streams from its catchment
area, where 83.3% of the catchment area is belongs to the Lore Lindu National
Park. However, generally the gentle slope eastern part and several small parts in
the western sites of the lake have been converted into agricultural land. Most of
the converted land has been turned into rice fields and cacao plantations. The
study area was divided to four i.e. north, east, south and west area.

4

Collecting specimen
Purposive sampling method was applied to 39 sampling sites (Fig. 1).
Samples were collected with tray net and hand net in July, August and November
2011. All specimens obtained were fixed in 96% ethanol. The ethanol was
changed after 24 hours with fresh 96% ethanol. The specimens are deposited in
Division of Zoology, Research Center for Biology, Indonesian Institute of
Sciences (LIPI) and Laboratory of Molecular, Division of Animal Function and
Behavior, Department of Biology, Bogor Agricultural University, Indonesia.
During collecting, we recorded environmental parameters i.e. 1) the
temperature of the water with a mercury thermometer, 2) the qualitative water
clarity, 3) the qualitative water current (slow current water to almost stagnant or
moderate flow), 4) substrate (sediment classification) based on Wentworth (1922),
5) presence or absence of water plant associated with the Caridina habitat, and 5)
the coordinate and altitude of sampling location, measured using GPS.
Environmental Data Analysis
The coordinate and altitude of each sampling site were overlaid to the map
of Lake Lindu system published by BAKOSURTANAL. The percentage of each
Caridina species found was measured by the amount of particularly species
caught to the total of Caridina caught.

Figure 2 Location of the Lake Lindu system in Sulawesi Island (right)
and overview of the sampling sites in the lake (left). Circle refer to the
sampling site

5
Morphometric Measurements for Taxonomical Analysis
Morphometric measurements were taken using a Nikon SMZ 800 stereomicroscope with an ocular micrometer. Drawings of chepalothorax were made
with a camera lucida mounted on the microscope while other cephalic and body
appendages were made with a microphotograph mounted on a Nikon Phase
Contrast 0.90 DRY light microscope. Drawings were then digitized and processed
with Adobe Photoshop CS3 extended.
Comparative data for Caridina linduensis is based on the original description
of Roux (1904), and for C. sarasinorum and C. schenkeli is based on the
published data of Cai and Wowor (2007) and von Rintelen and Cai (2009),
respectively. That for C. parvidentata and C. sulawesi is from Cai and Ng (2009).
Specimens examined for taxonomic work are deposited in Division of
Zoology (formerly known as Museum Zoologicum Bogoriense, MZB), Research
Center for Biology, Indonesian Institute of Sciences and ZRC, Zoological
Reference Collection, Raffles Museum of Biodiversity Research, National
University of Singapore. The abbreviation cl is used for carapace length measured
in mm from the post-orbital margin to the posterior median margin of the
carapace. Uwe is Kaili dialect of Central Sulawesi for stream while abbreviation P
is used for pulau, which translates to island in Indonesian.

6

4 RESULTS AND DISCUSSION
Results
Taxonomy of the Genus Caridina from Lake Lindu
Systematic account
Family Atyidae De Haan, 1849
Caridina H. Milne Edwards, 1837
Caridina linduensis Roux, 1904
(Figs. 3–4)
Caridina linduensis J. Roux 1904: 541, pl. 9, figs. 1–4 [type locality: Lake Lindu,
Sulawesi (Celebes), Indonesia].–Bouvier 1925: 224, figs. 497–503.–Chace
1997: 13.–von Rintelen et al. 2008: 2244.–De Grave and Fransen 2011: 276.
Material examined: Four males, cl 3.0–3.3 mm, 9 females, cl 3.6–4.4 mm,
6 ovigerous females, cl 3.8–4.2 mm (MZB Cru 3793), Uwe Rawa at the outlet of
Lake Lindu, about 50 m from the lake, 01°16’20.1”S 120°06’32.7”E, on leaf litter
and dead wood, coll. Annawaty and D. Wowor, 13 Nov 2011; 1 male, 1 ovigerous
female (ZRC), same data as MZB Cru 3793; 7 males, cl 2.7–3.4 mm, 4 females, cl
3.5–4.1 mm, 3 ovigerous females, cl 3.8–4.2 mm, 1 male, cl 3.5 mm, dissected
(MZB Cru 3794), southwest coast of P. Bola, 01°20’52.3”S 120°03’51.2”E, on
root of macrophytes, coll. Annawaty and D. Wowor, 17 Nov 2011; 5 males, cl
3.3–3.5 mm, 4 females, cl 3.8–4.7 mm, 5 ovigerous females, cl 4.0–4.9 mm, 1
female, cl 3.8 mm, dissected (MZB Cru 3795), at the mouth of Uwe Langko,
01°20’38.1”S 120°03’32.6”E, on root of macrophytes, coll. Annawaty and D.
Wowor, 17 Nov 2011; 1 female, cl 3.9 mm (MZB Cru 3796), at Tomado beach,
01°19’31.9”S 120°03’10.0”E, on root of macrophytes, coll. Annawaty and D.
Wowor, 15 Nov 2011.
Description: Rostrum slender, reaching near or slightly beyond end of third
segment of antennular peduncle, does not overreach end of scaphocerite, tip of
rostrum straight or slightly curved upwards, 0.5–0.7 times as long as carapace,
dorsal margin horizontal or slightly sinuous, dorsal with 7–12 (mode 9) teeth,
about one-third distal unarmed, no tooth behind orbital margin nor sub-apical
tooth, ventral with 0–6 (mode 2 or 3) teeth. Antennal spine short, situated below
inferior orbital angle. Pterygostomian margin rounded (Fig. 3A–C).
Eyes well developed, anterior end 0.6 times length of basal segment of
antennular peduncle. Antennular peduncle 0.6–0.8 times as long as carapace,
basal segment of antennular peduncle longer than second, third segment lengths,
second segment distinctly longer than third segment, anterolateral angle of basal
segment reaching 0.3–0.5 proximal of second segment of antennular peduncle
(Fig. 3E). Stylocerite reaching 0.7 of basal segment of antennular peduncle.

7
Scaphocerite 3.2 times as long as wide (Fig. 3D). Incisor process of mandible
ending in 16–18 irregular teeth, molar process truncated (Fig. 3F–G). Lower
lacinia of maxillula broadly rounded, upper lacinia elongated, with numerous
distinct teeth on inner margin, palp slender (Fig. 3H). Upper endite of maxilla

Figure 3 Caridina linduensis Roux, 1904.
A-C, cephalothorax and cephalic appendages, D. scaphocerite, E.
antennulla, F. mandible, G. incisor process of mandible H. maxillula, I.
maxilla, J. first maxilliped, K. second maxilliped, L. third maxilliped.
Scales: A-C = 1.0 mm; D, E, L = 0.5 mm; F, H-K = 0.2 mm; G = 0.1 mm.
(A, D-L: female, cl 3.8 mm, MZB Cru 3795; B: ovigerous female, cl 3.8
mm, egg size 0.9 x 0.6 mm, MZB Cru 3794; C: ovigerous female, cl 4.1
mm, egg size 0.9 x 0.6 mm, MZB Cru 3794)

8

Figure 4 Caridina linduensis Roux, 1904.
A. first pereiopod, B. second pereiopod, C. third pereiopod, D. dactylus
of third pereiopod, E. fifth pereiopod, F. dactylus of fifth pereiopod, G.
endopod of male first pereiopod, H. appendix masculina and appendix
interna of male second pleopod, I. distal portion of telson, J. telson, K.
diaeresis, L. preanal carina. Scales: A-C, E, J = 0.5 mm; G, H, L = 0.2
mm; D, F, I, K = 0.1 mm. (A-F, I-L: female, cl 3.8 mm, MZB Cru
3795; G-H: male, cl 3.5 mm, MZB Cru 3794)

9
subdivided, palp short, scaphognathite tapering posteriorly with numerous long,
curved setae at posterior end (Fig. 3I). Palp of first maxiliped ending in triangular
projection (Fig. 3G). Podobranch of second maxiliped reduced to lamina (Fig. 3K).
Third maxiliped reaching end of scaphocerite, with ultimate segment slightly
longer than penultimate segment (Fig. 3L).
Epipod present on first 2 pereiopods, reduced in size posteriorly, absent on
last 3 pereiopods. Chela and carpus of first pereiopod distinctly stouter, broader
than chela and carpus of second pereiopod. First pereiopod stout, reaching to
distal end of basal segment of antennular peduncle; merus 1.8–2.7 times as long
as wide, slightly shorter than carpus; carpus excavated anteriorly, 1.6–2.4 times as
long as wide, shorter than chela, chela 2.1–2.9 times as long as wide; movable
finger as long as or slightly longer than palm, finger tip round (Fig. 4A).
Second pereiopod reaching to distal end of third segment of antennular
peduncle; merus distinctly shorter than carpus, 4.4–5.8 times as long as wide;
carpus slender, 5.1–6.9 times as long as wide, 1.2–1.3 times length of chela; chela
2.5–3.7 times as long as wide; movable finger 1.2–1.8 times as long as palm,
finger tip round (Fig. 4B).
Third pereiopod slender, dactylus, half distal propodus overreach third
segment of antennular peduncle; propodus 10.4–13.0 times as long as wide, 4.1–
4.5 times as long as dactylus; dactylus 2.8–4.0 times as long as wide (terminal
spine included, without spines of flexor margin), terminating in 1 large claw with
5–9 (mode 5) accessory spines on flexor margin, reducing in size proximally (Fig.
4C–D). Sexual dimorphism present. Third, fourth pereiopods of male with
numerous hooked spinules on inner, outer margins of propodus, carpus,
respectively.
Fifth pereiopod slender, tip of dactylus reach end third segment of antennular
peduncle; propodus 10.8–13.7 times as long as wide, 2.5–3.3 times as long as
dactylus; dactylus 4.7–5.8 times as long as wide (terminal spine included, without
spines of flexor margin), terminating in 1 large claw, with 53–56 (mode 54)
accessory spinules on flexor margin (Fig. 4E–F).
Endopod of male first pleopod subrectangular, without appendix interna, 2.8
times as long as wide, 0.4 times length of exopod (Fig. 4G).
Appendix masculina of male second pleopod very slender, 0.5–0.6 times
length of endopod, with appendix interna 0.3–0.5 times length of appendix
masculina (Fig. 4H).
Sixth abdominal somite 0.5–0.6 times of carapace, 1.4–1.6 times as long as
fifth somite, 0.9 times as long as telson. Telson 2.8 times as long as wide, distal
margin rounded without posteromedian projection, with 4–6 (mode 5) pairs of
dorsal spinules, 1 pair of dorsolateral spinules; distal end with 3 or 4 (mode 3)
pairs of spine, lateral pair of spines distinctly longer than intermediate pairs (Fig.
4I–J). Preanal carina sub-rectangular, without spine (Fig. 4L).
Uropodal diaresis with 14–16 (mode 16) movable spinules (Fig. 4K).
Ovigerous females with 26–47 eggs (n = 3); Egg size 1.0–1.1 x 0.7 mm in
diameter (n = 40, eggs with eyes).
Habitat: Caridina linduensis is a soft substrate dweller on leaf litter, roots of
macrophytes and dead wood. It mainly occurs in the lake but can be also found in

10
the mouth of the only outlet stream with mud-sandy substrates with very slow
currents or almost stagnant water. The temperature of the habitat varies between
28.0° and 29.0°C.
Distribution: Caridina linduensis is found only in Lake Lindu and the mouth
of outlet stream. It is distributed in the south and southwestern of the lake, and
around the junction of the lake and Uwe Rawa at the north. A single specimen
was also caught at the west coast of the lake at Tomado village.
Remarks: Although we did not compare our specimens with the type
material of C. linduensis, but we have seen the excellent drawings of the type
material in the thesis of Cai (2004) which confirms our observations that our
collected specimens from the type locality is indeed C. linduensis. However, our
material has few differences from those examined by Roux (1904) in the higher
number of accessory spines on the flexor margin of the fifth pereiopods (53-56,
mode 54 vs. 50) and the ratio of the dactylus to the palm of the first pereiopod
(dactylus as long as or slightly longer than palm vs. dactylus slightly shorter than
palm). The differences might be an intraspecific variation within the population.
In this study, we observe that there are two characters which were not
mentioned by Roux (1904), i.e. the mandible and the presence of sexual
dimorphism in C. linduensis. The results show that the terminal of the incisor
process of the mandible is ending in many irregular sharp teeth, and the inner
margin of the propodus and the outer margin of the carpus of both third and fourth
pereiopods of males are covered by numerous claw-shape spinules.
With regard to the shape of the rostrum, the position of the antennal spine to
the inferior orbital angle, and the absence of the posteromedian projection of the
telson, C. linduensis resembles C. sarasinorum and C. schenkeli, two endemic
Caridina species from Lake Poso in Sulawesi. However, it can be separated from
both species by the absence of tooth behind the orbital margin (vs. 3–7 teeth in C.
sarasinorum, 2–5 teeth in C. schenkeli), the lower number of the ventral rostral
teeth (0–6 teeth vs. 8–17 teeth in C. sarasinorum, 9–13 teeth in C. schenkeli), the
smaller ratio of the second segment to the third segment of the antennular
peduncle (1.0–1.3 vs. 2.0 in C. sarasinorum, 1.6–2.0 in C. schenkeli), the smaller
ratio of the antennular peduncle length to the carapace length (0.6–0.8 times vs. as
long as in C. sarasinorum, 0.8–1.0 times in C. schenkeli), the relatively broader
scaphocerite (3.2 times as long as wide vs. 4.4 times in C. sarasinorum, 3.3–4.7
times in C. schenkeli), the more slender dactylus of the fifth pereiopod (4.7–5.8 as
long as wide vs. 3.0–3.9 in C. sarasinorum, 3.7–4.8 in C. schenkeli), the broader
telson (2.8 times as long as wide vs. 4.1 times in C. sarasinorum, 3.0–3.6 times in
C. schenkeli), the smaller ratio of the sixth abdominal somite to the fifth
abdominal somite (1.4–1.6 vs. 2.0 in C. sarasinorum, 1.9–2.0 in C. schenkeli), the
absence of preanal carina spine (vs. with spine in both C. sarasinorum and C.
schenkeli), and the higher number of the uropodal diaresis spinules (14–16 vs. 8–
10 in C. sarasinorum, 10–11 in C. schenkeli). Caridina linduensis can also be
separated from C. sarasinorum and C. schenkeli by the number of the accessory
spinules on the flexor margin of the fifth pereiopod (53–56 vs. 33–43 in C.
sarasinorum, 57–64 in C. schenkeli), and the different shape of the endopod of the
first pleopod of male (sub-rectangular in C. linduensis, vs. sub-triangular in C.
sarasinorum, elongated triangular in C. schenkeli).

11
Caridina linduensis also shares several characters with C. schenkeli which
can be used to distinguished both species from C. sarasinorum, i.e., by the
presence of epipod on the first two pereiopods (vs. present on the first pereiopod
and greatly reduced or absent on the second pereiopod in C. sarasinorum), and the
larger egg size (1.0–1.1 x 0.7 mm vs. 0.9–1.0 x 0.5–0.6 mm in C. sarasinorum).
Caridina linduensis can also be distinguished from C. schenkeli by the smaller
ratio of the appendix interna to the appendix masculina of the male second
pleopod (0.3–0.5 vs. 0.7–0.8 in C. schenkeli) and from C. sarasinorum by the
lower number of the dorsal rostral teeth (7–12 teeth vs. 12–19 teeth in C.
sarasinorum)
Caridina sp B
(Figs. 5–6)
Material examined: Holotype: ovigerous female (eggs with eyes), cl 4.7
mm (MZB Cru 3806), Uwe Wobo at Jambu-Jambu, Lake Lindu, 01°20’30.3”S
120°05’17.2”E, on gravel, and roots of fern and grass, coll. Annawaty and D.
Wowor, 21 Nov 2011. Paratypes: 11 females, cl 3.9–4.8 mm, 2 ovigerous females
4.2–4.9 mm (MZB Cru 3807), same data as holotype; 1 male, cl 3.5 mm, 1
ovigerous female, cl 4.4 mm (MZB Cru 3808), Uwe Poweroa, Lake Lindu,
01°20’05.6”S 120°05’55.2”E, on roots of macrophytes, coll. Annawaty and D.
Wowor, 21 Nov 2011; 12 females, cl 3.8–4.5 mm (MZB Cru 3809), at the mouth
of Air Dingin stream, Lake Lindu, 01°20’07.2’’S 120°05’52.2’’E, on roots of
macrophytes, coll. Annawaty and D. Wowor, 21 Nov 2011; 2 males, cl 3.2 mm, 1
female, cl 4.8 mm (MZB Cru 3810), west side of mouth of Uwe Kati, Lake Lindu,
01°20’54.4”S 120°04’51.0”E, on roots of macrophytes, coll. Annawaty and D.
Wowor, 18 Nov 2011; 2 females, cl 3.8 mm (MZB Cru 3811), at east side of
mouth of Uwe Kati, Lake Lindu, 01°20’54.4”S 120°04’51.0”E, on roots of
macrophytes, coll. Annawaty and D. Wowor, 16 Nov 2011; 26 males, cl 3.7–4.8
mm, 13 females, cl 4.0–5.3 mm, 5 ovigerous females, cl 4.8–5.4 mm, 1 male, cl
4.1 mm, dissected (MZB Cru 3812), Uwe Kati, Lake Lindu catchment,
01°21’54.5”S 120°05’35.2”E, on roots of macrophytes, coll. Annawaty and D.
Wowor, 21 Nov 2011; 1 male, 1 female, 1 ovigerous female (ZRC), same data as
MZB Cru 3812. Others: 29 juveniles, 8 males, 9 females, 1 ovigerous female
(MZB Cru 3974), Uwe Kati, Lake Lindu catchment, coll. Annawaty and D.
Wowor, 21 Nov 2011.
Description: Rostrum short, overreach end of basal segment to half
proximal of second segment of antennular peduncle, 0.4 times as long as carapace,
dorsal margin bent downwards above orbit with tip directed anteriorly, dorsal with
1–9 (mode 6) teeth, without tooth behind orbital margin, no sub-apical tooth,
ventral with 0–3 (mode 1) teeth. Antennal spine short, situated below inferior
orbital angle. Pterygostomian margin rounded (Fig. 5A).
Eyes well developed, anterior end 0.6 times length of basal segment of
antennular peduncle. Antennular peduncle 0.6–0.7 times as long as carapace,
basal segment of antennular peduncle longer than second, third segment lengths,
second segment distinctly longer than third segment, anterolateral angle of basal
segment reaching 0.3 second segment of antennular peduncle. Stylocerite reaching

12
0.7 basal segment of antennular peduncle (Fig. 5C). Scaphocerite 3.2 times as
long as wide (Fig. 5B).
Incisor process of mandible ending in 6 irregular teeth and 2 very fine
granulated rows between teeth, molar process straight (Fig. 5D–E). Lower lacinia
of maxillula broadly rounded, upper lacinia elongated, with numerous distinct
teeth on inner margin, palp slender (Fig. 5F). Upper endite of maxilla subdivided,
palp short, scaphognathite tapering posteriorly with numerous long, curved setae
at posterior end (Fig. 5G). Palp of first maxiliped ending in triangular projection
(Fig. 5H). Podobranch of second maxiliped reduced to a lamina (Fig. 5I). Third
maxiliped reaching end of scaphocerite with ultimate segment slightly longer than
penultimate segment (Fig. 5J).
Epipod present on first 2 pereiopods, reduced in size posteriorly, absent on
last 3 pereiopods. Chela and carpus of first pereiopod distinctly stouter, broader
than chela and carpus of second pereiopod.
First pereiopod stout, tip of chela not reaching basal segment of antennular
peduncle; merus 2.2–2.5 times as long as wide, slightly shorter than carpus;
carpus excavated anteriorly, distinctly shorter than chela, 1.7–2.5 times as long as
wide, chela 2.0–2.4 times as long as wide; movable finger slightly longer than
palm, finger tip round (Fig. 6A).
Second pereiopod reach end of third segment of antennular peduncle; merus
distinctly shorter than carpus, 4.1–4.8 times as long as wide; carpus slender, 1.3–
1.4 times as long as chela, 6.1–6.7 times as long as wide; chela 3.2–3.5 times as
long as wide; movable finger 1.4–1.6 times as long as palm, finger tip round (Fig.
6B).
Third pereiopod slender, dactylus, half distal propodus overreach third
segment of antennular peduncle; propodus 10.0–10.6 times as long as wide, 3.6–
4.2 times as long as dactylus; dactylus 3.3–4.2 times as long as wide (terminal
spine included, without spines of flexor margin), terminating in 1 large claw with
7–10 (mode 7) accessory spines on flexor margin, reducing in size proximally
(Fig. 6C–D). Sexual dimorphism present. Third and fourth pereiopods of male
with numerous hooked spinules on inner, outer margins of propodus, carpus
respectively.
Fifth pereiopod slender, one-third distal of dactylus overreach third segment
of antennular peduncle; propodus 11.5–13.7 times as long as wide, 2.7–3.2 times
as long as dactylus; dactylus 5.5–6.0 times as long as wide (terminal spine
included, without spines of flexor margin), terminating in 1 large claw with 63–80
accessory spinules on flexor margin (Fig. 6E–F).
Endopod of male first pleopod subrectangular, without appendix interna, 3.2
times as long as wide, 0.4–0.5 times length of exopod (Fig. 6G).
Appendix masculina of male second pleopod very slender, 0.5–0.6 times
length of endopod, with appendix interna 0.4–0.5 times length of appendix
masculina (Fig. 6H).
Sixth abdominal somite 0.5 times as long as carapace, 1.7–1.8 times as long
as fifth somite, 0.9 times as long as telson. Telson 2.7 times as long as wide, distal
margin rounded without posteromedian projection, with 4 pairs of dorsal spinules,
1 pair of dorsolateral spinules; distal end with 3–6 (mode 3) pairs of spine, lateral
pair of spines distinctly longer than intermediate pairs (Fig. 6I–J). Preanal carina
subtriangular, without spine (Fig. 6L).

13

Figure 5 Caridina sp B.
A cephalothorax and cephalic appendages, B. scaphocerite, C. antennulla,
D. mandible, E. incisor process of mandible F. maxillula, G. maxilla, H.
first maxilliped, I. second maxilliped, J. third maxilliped. Scales: A = 1.0
mm; B, C, J = 0.5 mm; D, F-I = 0.2 mm; E = 0.1 mm. (A: paratype,
ovigerous female, cl 4.5 mm, MZB Cru 3807; B-J: paratype, male, cl 4.1
mm, MZB Cru 3812)

14

Figure 6 Line drawing of Caridina sp B.
A. first pereiopod, B. second pereiopod, C. third pereiopod, D. dactylus
of third pereiopod, E. fifth pereiopod, F. dactylus of fifth pereiopod, G.
endopod of male first pereiopod, H. appendix masculina and appendix
interna of male second pleopod, I. distal portion of telson, J. telson, K.
diaeresis, L. preanal carina. Scales: A-C, E, J = 0.5 mm; G, H, L = 0.2
mm; D, F, I, K = 0.1 mm. (A-L: paratype, male, cl 4.1 mm, MZB 3812)

15
Uropodal diaresis with 13–15 (mode 15) movable spinules (Fig. 6K).
Ovigerous females with with 39–40 eggs (n = 2); Egg size 1.1–1.2 x 0.7–0.8
mm in diameter (n= 40, eggs with eyes).
Habitat: Caridina sp B is a soft substrate dweller on roots of macrophytes
(water grass and ferns). This species mainly occurs at the edge of inlet streams
and the edge of the lake that have muddy-sandy substrates. It was found in
streams of 0.5 to 3 m depth, with murky and slow running water that may be
almost stagnant. The temperature of the habitat varies from 22.0° to 28.5°C.
Distribution: Caridina sp B is endemic to Lake Lindu and the catchment
area. So far, it was found only at the relatively flat southeastern shore of the lake
and Uwe Kati. This species has not been encountered in the north and the west
coasts of the lake.
Etymology: The species is named after its main habitat, a local name for
water grass, where this species mainly occur clinging around. The name is used as
a noun in apposition.
Remarks: With regard to the shape of the rostrum, the absence of a tooth
behind the orbital margin, the position of the antennal spine to the inferior orbital
angle, the absence of a median projection at the terminal margin of the telson, the
absence of a preanal carina spine, and the absence of an appendix interna at the
terminal margin of the sub-rectangular endopod of the first pleopod of male,
Caridina sp B is similar to C. linduensis and C. parvidentata Roux, 1904.
However, the new species can be differentiated from C. linduensis by the shorter
rostrum (ratio of rostrum length to carapace length 0.4 vs. 0.5–0.7 in C.
linduensis), the less number of dorsal rostral teeth (1–9, mode 6 vs. 7–12, mode 9
in C. linduensis), the fewer number of ventral rostral teeth (0–3, mode 1 vs. 0–6,
mode 3 in C. linduensis), the fewer number of irregular teeth of the incisor
process of the mandible (6 vs. 16–18 irregular teeth in C. linduensis), and the
larger number of accessory spinules on the flexor margin of the dactylus of the
fifth pereiopod (63–80 vs. 53–56, mode 54).
Caridina sp B can also be distinguished from C. parvidentata by the
position of the antennal spine (below inferior orbital angle vs. fused with inferior
orbital angle in C. parvidentata), the brouder scaphocerite (3.2 times as long as
wide vs. 3.8 times in C. parvidentata), the presence of epipod on the first two
pereiopods (vs. on the first three pereiopods in C. parvidentata), the stouter chela
of the second pereiopod (3.2–3.5 times as long as wide vs. 3.8 times in C.
parvidentata), the more slender dactylus of the fifth pereiopod (5.5–6.0 times as
long as wide vs. 4.1 times in C. parvidentata), the lower ratio of appendix
masculina to endopod of male second pleopod (0.5–0.6 times vs. 0.8 times in C.
parvidentata), the proportionately broader telson (2.7 times as long as wide vs. 3.0
times in C. parvidentata), the smaller ratio of sixth abdominal somite to telson
length (0.9 vs. as long as in C. parvidentata), and the relatively larger egg size
(1.1–1.2 x 0.7–0.8 mm vs. 0.9 x 0.6 mm in C. parvidentata). In addition, C. sp B
can be distinguished from both C. linduensis and C. parvidentata by having more
accessory spinules on the flexor margin of the fifth pereiopod (63–80 vs. 53–56 in
C. linduensis, 61 in C. parvidentata).

16
Caridina sp A
(Figs. 7–8)
Material examined: Holotype: ovigerous female (eggs with eyes), cl 5.2
mm (MZB Cru 3797), Uwe Kaongko, Lake Lindu catchment, 01°18’57.2”S
120°03’08.4”E, on gravel and roots of macrophytes, coll. Annawaty and D.
Wowor, 15 Nov 2011. Paratypes: 25 males, cl 4.7–6.0 mm, 46 females, cl 4.4–5.6
mm, 14 ovigerous females, 5.4–6.8 mm (MZB Cru 3798), same data as holotype;
1 male, 1 female, 1 ovigerous female (ZRC), same data as holotype; 13 males, cl
3.4–4.5 mm, 17 females, cl 4.6–6.7 mm, 11 ovigerous females, cl 5.1–5.7 mm
(MZB Cru 3799), Uwe Kumo, Lake Lindu catchment, 01°16’46.3”S
120°04’05.8”E, on gravel, boulders, roots of macrophytes, coll. Annawaty and D.
Wowor, 20 Nov 2011; 20 males, cl 4.4–5.5 mm, 11 females, cl 5.1–6.0 mm, 28
ovigerous females, cl 5.0–6.3 mm (MZB Cru 3800), Uwe Tokaroru, Lake Lindu
catchment, 01°17’08.7”S 120°07’15.5”E, on gravel and water moss, coll.
Annawaty and D. Wowor, 13 Nov 2011; 17 males, cl 3.9–5.1 mm, 22 females cl
5.0–6.2 mm, 18 ovigerous females, cl 5.1–6.0 mm, 1 male, cl 4.8 mm, dissected
(MZB Cru 3801), Uwe Lembosa, Lake Lindu catchment, 01°19’44.6”S
120°09’03.2”E, on gravel and roots of macrophytes, coll. Annawaty and D.
Wowor, 14 Nov 2011; 12 females, cl 4.4–5.7 mm, 3 ovigerous females, cl 4.8–5.3
mm (MZB Cru 3802), Kana tributary, Lake Lindu catchment, at about 100 m
north of 01°19’50.6”S 120°02’56.8”E, on gravel and leaf litter, coll. Annawaty
and D. Wowor, 17 Nov 2011; 7 males, cl 3.5–4.7 mm, 6 females, cl 3.9–4.6 mm,
3 ovigerous females, cl 4.6–5.9 (