# Revision of the species of Caridina (Decapoda: Atyidae) from the Western Indian Ocean islands with description of a new species

Valentin de Mazancourt, Marion Mennesson, Gérard Marquet, Philippe Keith

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Valentin de Mazancourt, Marion Mennesson, Gérard Marquet, Philippe Keith. Revision of the species of Caridina (Decapoda: Atyidae) from the Western Indian Ocean islands with description of a new species. European Journal of Taxonomy, 2025, 1007, pp.57-86. 10.5852/ejt.2025.1007.2995. hal-05214338

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Research article

urn:lsid:zoobank.org:pub:8037ABB8-4C81-4340-A8F4-42D1BFE638D4

# Revision of the species of *Caridina* (Decapoda: Atyidae) from the Western Indian Ocean islands with description of a new species

Valentin DE MAZANCOURT¹,* , Marion MENNESSON<fnref n="2" /> ,
Gérard MARQUET<fnref n="3" /> тан & Philippe KEITH<fnref n="4" /> тан

¹,<fnref n="2" />,<fnref n="4" /> Laboratoire de Biologie des Organismes et Écosystèmes Aquatiques (UMR 8067 BOREA), Muséum national d'Histoire naturelle, CNRS-8067, IRD-207, Sorbonne Université, UCN, UA, Equipe Biodiversité, Plasticité, Adaptation et Conservation, 57 rue Cuvier, CP26, 75005 Paris, France.

³ 96 rue de Richelieu, 75002 Paris, France.

**Abstract.** We provide an updated checklist and an identification key of the species of *Caridina* living in freshwater habitats of the Western Indian Ocean islands, including the Seychelles, the Comoros, Mayotte, La Réunion and Mauritius. We list six species including one new for science: *Caridina mauritii*, *Caridina spathulirostris*, *Caridina similis*, *Caridina natalensis*, *Caridina typus* and *Caridina henriettae* sp. nov. Molecular and morphological evidence led us to consider *Caridina richtersii* as a junior synonym of *C. spathulirostris*. Due to different life histories, some species are endemic to their island or archipelago, like *C. mauritii* and *C. spathulirostris*, endemic to Mauritius, or *C. similis* endemic to the Seychelles, whereas others have a wider distribution in the Western Indian Ocean like *C. natalensis* or *C. henriettae* sp. nov. or have an Indo-West Pacific distribution like *C. typus*.

**Keywords.** Integrative taxonomy, morphology, 16S, amphidromy, endemic.

de Mazancourt V., Mennesson M., Marquet G. & Keith P. 2025. Revision of the species of *Caridina* (Decapoda: Atyidae) from the Western Indian Ocean islands with description of a new species. *European Journal of Taxonomy* 1007: 57–86. https://doi.org/10.5852/ejt.2025.1007.2995

## Introduction

Comoros, Seychelles and Mascarenes are three archipelagos in the Southwest of the Indian Ocean (Fig. 1). The Comoros archipelago, equidistant from the African continent and Madagascar (around 300 km), includes the islands of Grande Comore, Anjouan, Mohéli and Mayotte. The Mascarene archipelago includes three main islands: La Réunion, Mauritius and Rodrigues. The Seychelles archipelago, comprising 115 islets, includes four main islands with small rivers or swamps: La Digue, Mahé, Praslin and Silhouette.

<footnote>*Corresponding author: valentin.demazancourt_ext@mnhn.fr</footnote>

<footnote n="2">²Email: marion.mennesson@mnhn.fr</footnote>

<footnote n="3">³Email: gmarquet@neuf.fr</footnote>

<footnote n="4">⁴Email: philippe.keith@mnhn.fr</footnote>
Between 1996 and 2013, rivers of several South West Indian Ocean (SWIO) islands were surveyed for atyid freshwater shrimps (Crustacea: Decapoda: Caridea) during field trips conducted by the Muséum national d'Histoire naturelle (MNHN) in the Mascarene archipelago (La Réunion, Mauritius and Rodrigues; see Keith *et al.* 1999, 2006), in Comoros (Anjouan, Mohéli) and Mayotte (see Keith *et al.* 2006) and in Seychelles (La Digue, Mahé, Praslin and Silhouette; see Keith *et al.* 2006). Keith *et al.* 2006 reported seven species of *Caridina* in the SWIO islands, namely *Caridina longirostris* H. Milne-Edward, 1837, *C. mauritii* Bouvier, 1912, *C. richtersi* Thallwitz, 1891, *C. serratirostris* De Man 1892, *C. similis* Bouvier, 1904, *C. spathulirostris* Richters, 1880 and *C. typus* H. Milne-Edwards, 1837. More recently, in La Réunion and Mayotte, ‘Ocea Consult’ conducted yearly inventories to monitor the aquatic communities. In Seychelles, since 2020, the MNHN and the NGO Gaea conduct baseline surveys and annual biodiversity inventories of fish, micro- and macro-invertebrates in 15 rivers including the collection of genetic materials. These studies led to the discoveries of new occurrences of species at several sites and have improved our knowledge on the freshwater shrimps in the SWIO.

The purpose of this study is to provide an updated checklist of the species of *Caridina* from the SWIO by using an integrative taxonomic approach to assess the validity of the species listed previously and check their geographical distributions in our study area.

<figure><img src="image_2.png" /><figcaption>**Fig. 1.** Map of the South West Indian Ocean with its main archipelagos (dashed lines) and islands where *Caridina* H. Milne Edwards, 1837 were found (modified from Mulochau *et al.* 2014).</figcaption></figure>
**Material and methods**

**Sampling**

Specimens from SWIO were collected by electrofishing (portable Dekka 3000 electric device, Germany or SAMUS 1000, Poland: http://www.electro-fisher.net/) or by hand. All material preserved in 75-95% ethanol has been deposited in the collections of the Muséum national d'Histoire naturelle in Paris (MNHN). Specimens were measured electronically under a stereo microscope using LAS X software (Leica).

**DNA extraction, amplification and sequencing**

DNA was isolated from abdominal tissues using the semi-automatic Eppendorf ep-Motion 5075 robot following the NucleoSpin 96 Tissue Core (Macherey-Nagel) protocol. Fragments of the mitochondrial marker 16S rRNA (~ 510 bp) were amplified using the pair of primers 16Sar-Lmod (TAC TTC TGC CTG TTT ATC AAAAA)/16Sbmod (GGT CTG AAC TCA AAT CAT GTAAA) (de Mazancourt et al. 2019a). DNA amplification was performed in 20 µl PCR reactions, containing approximately 3 ng of template DNA, 2.5 mM MgCl₂, 0.26 mM of each nucleotide, 0.3 µM of each primer, 5% DMSO, 1 ng of BSA and 1.5 units of QBIOTAQ polymerase (MP Biomedicals). Amplification products were generated by an initial denaturation step of 4 min at 94°C followed by 35 cycles of denaturation at 94°C for 30 s, annealing at 52°C (16S) for 40 s, extension at 72°C for 60 s and a final extension step at 72°C for 7 min. PCR products were sequenced using the same primers and in both directions to insure the accuracy of base calls. Chromatograms were viewed and edited using Geneious ver. 8 software (http://www.geneious.com/ (Kearse et al. 2012). All sequences are deposited in GenBank (Numbers PQ963381 to PQ963410, see Table 1).

**Molecular analyses**

DNA sequences were aligned using MEGA11 software (Tamura et al. 2021) with Muscle algorithm (Edgar 2004). Using Bayesian information criterion in jModelTest (Guindon & Gascuel 2003; Darriba et al. 2012) we retained the GTR + G + I model. Best-scoring ML trees were estimated using RAxML HPC2 ver. 8.2.10 (Stamatakis 2006) and best-scoring Bayesian Inference (BI) trees were estimated using MrBayes ver. 3.2.7 (Ronquist & Huelsenbeck 2003), both methods implemented in CIPRES with the previously determined model, running for 10 000 000 generations, a sampling frequency of 2000 and a burn in of 10%. Support for nodes was determined using posterior probabilities calculated by MrBayes implemented in the Cyber Infrastructure for Phylogenetic Research (CIPRES) portal ver. 3.1. (https://www.phylo.org; Miller et al. 2010). One hundred independent searches, each starting from distinct random trees, were conducted. Robustness of the nodes was assessed using non-parametric bootstrapping (Felsenstein 1985) with 1000 bootstrap replicates. For the analysis, we generated 30 new 16S sequences from specimens collected during the authors' field trips, to which were added 24 sequences retrieved from GenBank for species occurring in the SWIO and/or closely related species to support the morphological comparisons (Table 1).

**Morphological study**

The rostrum, the cephalothorax, the pereiopods 1-3 and 5 and the abdomen were observed using a stereo microscope. The proportions of the various joints of the appendages were measured using microphotographs and the LAS X software (Leica). Drawings were made using the “Digital Inking” method (Coleman 2003, 2006) by tracing vectorial paths on high-resolution photographs using a vector graphics software.
<table><caption>**Table 1 (continued on next page). List of specimens included in the molecular analysis.**</caption><thead><tr><th>Species</th><th>Locality</th><th>Collection number</th><th>DNA voucher</th><th>Type status</th><th>16S sequence</th><th>Reference</th></tr></thead><tbody><tr><td rowspan="4">Atyoida serrata<br>Bate, 1888</td><td rowspan="4">La Réunion</td><td></td><td>CA2948</td><td></td><td>PQ963394</td><td>this study</td></tr><tr><td></td><td>CA3042</td><td></td><td>PQ963401</td><td>this study</td></tr><tr><td></td><td>CA3043</td><td></td><td>PQ963402</td><td>this study</td></tr><tr><td></td><td>CA3064</td><td></td><td>PQ963409</td><td>this study</td></tr><tr><td rowspan="3">Caridina spathulirostris<br>Richters, 1880</td><td rowspan="3">Mauritius</td><td>MNHN-2025-125</td><td>CA2165</td><td></td><td>PQ963389</td><td>this study</td></tr><tr><td>MNHN-2025-126</td><td>CA2166</td><td></td><td>PQ963390</td><td>this study</td></tr><tr><td>MNHN-2025-127</td><td>CA2167</td><td></td><td>PQ963391</td><td>this study</td></tr><tr><td rowspan="4">Caridina similis<br>Bouvier, 1904</td><td rowspan="4">Seychelles</td><td>MNHN-2025-130</td><td>CA1854</td><td></td><td>PQ963384</td><td>this study</td></tr><tr><td>MNHN-2025-131</td><td>CA1855</td><td></td><td>PQ963385</td><td>this study</td></tr><tr><td>MNHN-2025-132</td><td>CA1856</td><td></td><td>PQ963386</td><td>this study</td></tr><tr><td>MNHN-2025-133</td><td>CA1857</td><td></td><td>PQ963387</td><td>this study</td></tr><tr><td rowspan="12">Caridina natalensis<br>De Man, 1908</td><td rowspan="2">South Africa</td><td>MNHN-IU-2018-63</td><td>CA2080</td><td></td><td>OR088398</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td>MNHN-IU-2018-61</td><td>CA2079</td><td></td><td>MH644428</td><td>de Mazancourt<br>et al. 2019a</td></tr><tr><td rowspan="2">Madagascar</td><td></td><td>CA1074</td><td></td><td>OR088348</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td></td><td>CA1075</td><td></td><td>OR088349</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td rowspan="2">Mayotte</td><td></td><td>CA1077</td><td></td><td>OR088350</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td></td><td>CA1079</td><td></td><td>OR088351</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td rowspan="2">Comoros</td><td>MNHN-IU-2018-78</td><td>CA1343</td><td></td><td>MH644416</td><td>de Mazancourt<br>et al. 2019a</td></tr><tr><td>MNHN-IU-2018-77</td><td>CA1344</td><td></td><td>MH644417</td><td>de Mazancourt<br>et al. 2019a</td></tr><tr><td rowspan="2">Seychelles</td><td></td><td>CA1116</td><td></td><td>OR088365</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td></td><td>CA1117</td><td></td><td>OR088366</td><td>de Mazancourt<br>et al., 2023</td></tr><tr><td rowspan="4">Mauritius</td><td>MNHN-IU-2017-1451</td><td>CA2157</td><td></td><td>OR088418</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td>MNHN-IU-2017-1452</td><td>CA2158</td><td></td><td>OR088419</td><td>de Mazancourt<br>et al., 2023</td></tr><tr><td>MNHN-IU-2017-1453</td><td>CA2159</td><td></td><td>OR088420</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td>MNHN-IU-2017-1454</td><td>CA2160</td><td></td><td>OR088421</td><td>de Mazancourt<br>et al. 2023</td></tr><tr><td rowspan="2">Solomon</td><td>MNHN-IU-2018-2927</td><td>CA1351</td><td></td><td>MT303939</td><td>de Mazancourt<br>et al. 2020</td></tr><tr><td>MNHN-IU-2018-2931</td><td>CA1523</td><td></td><td>MT303940</td><td>de Mazancourt<br>et al. 2020</td></tr><tr><td rowspan="3">Caridina serratirostris<br>De Man, 1892</td><td rowspan="3">Seychelles</td><td>MNHN-2025-156</td><td>CA1491</td><td>paratype</td><td>PQ963382</td><td>this study</td></tr><tr><td>MNHN-2025-157</td><td>CA1492</td><td>paratype</td><td>PQ963383</td><td>this study</td></tr><tr><td>MNHN-IU-2015-1932</td><td>CA2026</td><td>paratype</td><td>PQ963388</td><td>this study</td></tr></tbody></table>
<table><caption>**Table 1 (continued).**</caption><thead><tr><th>Species</th><th>Locality</th><th>Collection number</th><th>DNA voucher</th><th>Type status</th><th>16S sequence</th><th>Reference</th></tr></thead><tbody><tr><td rowspan="15">Caridina typus<br>H. Milne Edwards, 1837</td><td rowspan="15">La Réunion</td><td>MNHN-2025-154</td><td>CA2998</td><td>paratype</td><td>PQ963396</td><td>this study</td></tr><tr><td>MNHN-2025-155</td><td>CA2999</td><td>paratype</td><td>PQ963397</td><td>this study</td></tr><tr><td>MNHN-2025-151</td><td>CA3000</td><td>paratype</td><td>PQ963398</td><td>this study</td></tr><tr><td>MNHN-2025-143</td><td>CA2417</td><td>paratype</td><td>PQ963392</td><td>this study</td></tr><tr><td>MNHN-2025-144</td><td>CA2418</td><td>paratype</td><td>PQ963393</td><td>this study</td></tr><tr><td>MNHN-2025-145</td><td>CA2949</td><td>paratype</td><td>PQ963395</td><td>this study</td></tr><tr><td>MNHN-2025-148</td><td>CA3065</td><td>paratype</td><td>PQ963410</td><td>this study</td></tr><tr><td>MNHN-2025-146</td><td>CA3038</td><td>paratype</td><td>PQ963400</td><td>this study</td></tr><tr><td>MNHN-2025-147</td><td>CA3061</td><td>paratype</td><td>PQ963407</td><td>this study</td></tr><tr><td>OUMNH.ZC-2010-22-0001</td><td></td><td></td><td>KY069380</td><td>Bernardes<br>et al. 2017</td></tr><tr><td>OUMNH.ZC-2010-22-0003</td><td></td><td></td><td>KY069381</td><td>Bernardes<br>et al. 2017</td></tr><tr><td>OUMNH.ZC-2010-22-0007</td><td></td><td></td><td>KY069382</td><td>Bernardes<br>et al. 2017</td></tr><tr><td rowspan="6">La Réunion</td><td>MNHN-2025-135</td><td>CA3054</td><td></td><td>PQ963403</td><td>this study</td></tr><tr><td>MNHN-2025-136</td><td>CA3058</td><td></td><td>PQ963404</td><td>this study</td></tr><tr><td>MNHN-2025-137</td><td>CA3059</td><td></td><td>PQ963405</td><td>this study</td></tr><tr><td>MNHN-2025-138</td><td>CA3060</td><td></td><td>PQ963406</td><td>this study</td></tr><tr><td>MNHN-2025-139</td><td>CA3063</td><td></td><td>PQ963408</td><td>this study</td></tr><tr><td>ZMBunid1252</td><td></td><td></td><td>KY069434</td><td>Bernardes<br>et al., 2017</td></tr><tr><td>Australia</td><td>AM-P21882</td><td></td><td></td><td>DQ478562</td><td>Page et al. 2007</td></tr><tr><td>Vanuatu</td><td>GU-721</td><td></td><td></td><td>DQ478563</td><td>Page et al. 2007</td></tr><tr><td rowspan="2">Madagascar</td><td>MNHN-IU-2018-2829</td><td>CA1038</td><td></td><td>MT303907</td><td>de Mazancourt<br>et al. 2020</td></tr><tr><td>MNHN-IU-2018-2831</td><td>CA1039</td><td></td><td>PQ963381</td><td>this study</td></tr><tr><td>South Africa</td><td>MNHN-IU-2018-2844</td><td>CA2090</td><td></td><td>MT303906</td><td>de Mazancourt<br>et al. 2020</td></tr><tr><td>Mayotte</td><td>MNHN-IU-2025-134</td><td>CA3027</td><td></td><td>PQ963399</td><td>this study</td></tr></tbody></table>

# Results

## Molecular analyses

Phylogenetic analyses of the 16S marker (Fig. 2) allowed to recognize 7 different clades including 6 from the SWIO, all of them being strongly supported in both analyses (Bayesian PP > 0.95 and ML bootstraps > 92). Three species complexes can be identified, all with very strong statistical support values (PP = 1; B = 100).

## Morphological study

Measures and observations made on the specimens allowed us to confirm the molecular results in recognizing the clades as five different species of *Caridina* in the SWIO, including one new for science (see hereafter). We couldn't get any sequence for *C. mauritii* due to the lack of fresh material, but its morphology is different enough to still be considered distinct from the other species. A key is provided at the end of the paper.
## Taxonomy

Class Malacostraca Latreille, 1802

Order Decapoda Latreille, 1802

Family Atyidae De Haan, 1849

Genus *Caridina* H. Milne Edwards, 1837

*Caridina nilotica* species group

## Diagnosis

Slender morphology, with a mostly long rostrum (longer than the antennular peduncle) but variable, the antennal spine ventral to the inferior orbital angle, a long antennular peduncle (subequal to carapace length), segments of walking legs slender, a typical dorsal hump over the third abdominal somite, a long sixth abdominal somite (always more than half of carapace length), a small pre-anal carina sometimes bearing an acute spine, a moderate number of spinules on the uropodal diaeresis (< 15), fewer, medium to short and terminal setae on the telson and a subtriangular endopod of the first male pleopod with or without an appendix on the subdistal outer margin or even placed at the distal end. An oblique red band on the cephalothorax is very characteristic.

*Caridina mauritii* Bouvier, 1912

*Caridina Mauritii* Bouvier, 1912a: 298.

*Caridina Mauritii* – Bouvier 1913a: 464 (key); 1914: 699; 1925: 210, figs 464–469.

*Caridina mauritii* – Costa 1980: 690. — Keith *et al*. 2006: 52.

<figure><img src="image_3.png" /><figcaption>**Fig. 2.** Bayesian inference phylogenetic tree obtained from the 16S marker. Numbers above branches represent Bayesian posterior probabilities, numbers under branches are Maximum Likelihood bootstraps. Letters indicate clades recognized as valid species and colored boxes are species complexes.</figcaption></figure>
**Type locality**

Near Port-Louis, Mauritius.

**Material examined**

No material examined.

**Distribution**

Endemic to Mauritius.

**Habitat**

Freshwater.

**Remarks**

This small species seems rare and a DNA barcode sequence would be needed to complete our sampling.

*Caridina natalensis* De Man, 1908
Figs 2D, 5B

*Caridina nilotica* var. *natalensis* De Man, 1908: 262, pl. 20 fig. 3.

*Caridina longirostris* — Richters 1880: 162–163. — Holthuis 1965: 20–23, fig. 6. — Keith *et al.* 2006: 50. — Bouchard *et al.* 2013: 9.

*Caridina wycki* — Ortmann 1894a: 11.1894b: 405 (part: specimens from East Africa and Dar-es-Salaam) (incorrect spelling).

*Caridina wyckii* — Weber 1897: 168 (part). — Richters 1880: 386 (part). — Lenz 1905: 385.

*Caridina nilotica* var. *brachydactyla* — Lenz 1910: 568. — Bouvier 1925: 155, figs 321–322. — Roux 1929: 303.

*Caridina nilotica* var. *gracilipes* — Lenz 1910: 568.

*Caridina nilotica* var. *natalensis* — Lenz 1912: 5. — Bouvier 1925: 154, fig. 320. — Barnard 1950: 655 (key). — Richard & Clark 2010: 317. — Klotz & De Grave 2015: 12.

*Caridina brachydactyla* *brachydactyla* — Costa 1980: 673.

*Caridina brachydactyla* — Bouchard *et al.* 2013: 9.

*Caridina natalensis* — de Mazancourt *et al.* 2019a: 379, figs 1–2, 3a–d; 2019b: 166, figs 2–5.

*Caridina umtatensis* — Wood *et al.* 2019: 205, fig. 3.

Not *Caridina africana* f. *natalensis* Bouvier, 1925: 214 (= *Elephantis jaggeri*).

Not *Caridina natalensis* — Richard & Clark 2009: 22, figs 7–8. — Castelin *et al.* 2013: 577 (= *Elephantis jaggeri*).

**Type locality**

Umgeni River, Durban, Natal, South Africa.

**Material examined**

See de Mazancourt *et al.* 2019a.

**Distribution**

Comoros (Anjouan, Mohéli), Mayotte, Seychelles (La Digue, Mahé, Praslin and Silhouette), Mauritius, La Réunion(?), Madagascar, South Africa, Tanzania and Kenya.
**Habitat**

Freshwater in the lower course of rivers.

**Remarks**

This species is easily identified by its rostrum rather constantly long, reaching well beyond scaphocerite.

*Caridina spathulirostris* Richters, 1880

Figs 2B, 5C

*Caridina serrata* Richters, 1880: 163, pl. 17 figs 24–27 (junior homonym of *Caridina serrata* Stimpson, 1860).

*Caridina spathulirostris* Richters, 1880: 163, pl. 17 fig. 28.

*Caridina spatulirostris* f. rectirostris Bouvier, 1925: 197.

*Caridina spatulirostris* f. curvirostris Bouvier, 1925: 197.

*Caridina richtersii* Thallwitz, 1891: 27 (nomen novum for *Caridina serrata* Richters, 1880).

*Caridina Richtersi* f. typica Bouvier, 1912c: 165; 1925: 203.

*Caridina apiocheles* Bouvier, 1904: 134.

*Caridina apiocheles* mutation *Edwardsi* Bouvier, 1904: 134.

*Caridina spathulirostris* – Ortmann 1894b: 406 (key). — Bouvier 1905: 93 (key); 1912a: 291; 1913a: 464 (key); 1925: 197, figs 416–422. — Costa 1980: 691. — Keith *et al.* 2006: 60.

*Caridina richtersi* – Ortmann 1894b: 402 (key), 404. — Edmondson 1929: 28. — Holthuis 1954: 3. — Carpenter 1978: 350. — Costa 1980: 690. — Keith *et al.* 2006: 54 (incorrect spelling).

*Caridina Richtersi* – Bouvier 1904: 134; 1905: 86; 1912a 293; 1912b: 919; 1912c: 163; 1913a: 464 (key); 1913b: 184; 1914: 699; 1925: 201, figs 432–457. — Bouvier & d’Emmerez de Charmoy 1919: 317. — Wolterock 1937a: 246 (incorrect spelling).

*Caridina apiocheles* – Bouvier 1905: 69, 86, fig. 3. — Bordage 1909: 108, fig. 6.

*Caridina apiocheles* mutation *Edwardsi* – Bouvier 1905: 69, 87, 103, fig. 3; 1912c: 166. — Bordage 1909: 108, fig. 6.

*Caridina Richtersi* f. *apiocheles* – Bouvier 1912a: 295; 1912c: 165; 1925: 203.

*Ortmannia Edwardsi* – Bouvier 1912 a: 296; 1912c: 166; 1912d: 691, fig. 3; 1913a: 459 (key); 1914: 700; 1925: 265, figs 445–447, 449, 452–453, 598–605. — Bouvier & d’Emmerez de Charmoy 1919: 317. — Wolterock 1937b: 322.

*Caridina richtersi* var. *apiocheles* – Bouvier 1925: 205. — Costa 1980: 690.

*Ortmannia edwardsi* – Edmondson 1929: 28. — Holthuis 1954: 3. — Costa 1980: 699.

*Caridina richtersi* var. *typica* – Costa 1980: 690.

**Type locality**

Pamplemousses Botanical Garden, Mauritius.

**Material examined**

*Caridina spathulirostris* Richters, 1880

MAURITIUS • 1 ♀ ovig. (cl 3.9 mm), 1 ♀ (cl 5.3 mm); quartier militaire, rivière importante; 20°17.260' S, 57°35.948' E; 390 m a.s.l.; 10 Dec. 2005; C.R. Turner leg.; MNHN-Na 772; MNHN • 1 ♀ (cl 5.8 mm); same data as for preceding; DNA voucher: CA2165; MNHN-2025-125; MNHN.
*Caridina richtersii* Thallwitz, 1891

MAURITIUS • 1 ♀ ovig. (cl 5.8 mm); quartier militaire, rivière importante; 20°17.260' S, 57°35.948' E; 390 m a.s.l.; 10 Dec. 2005; C.R. Turner leg.; DNA voucher: CA2166; MNHN-2025-126; MNHN • 1 ♂ (cl 6.5 mm); same data as for preceding; DNA voucher: CA2167; MNHN-2025-127; MNHN.

**Distribution**

Endemic to Mauritius.

**Habitat**

Freshwater in rivers and ponds.

**Remarks**

Our genetic study shows that *Caridina richtersi* Thallwitz, 1891 and *C. spathulirostris* Richters, 1880 are conspecific and we consider the former to be a junior synonym of *C. spathulirostris*. The specimens of this species are polymorphic in the shape of the rostrum and the chelipeds, which led to the description of numerous forms like *C. spathulirostris* f. rectirostris, *C. spathulirostris* f. curvirostris, *C. richtersi* f. typica, *C. richertsi* f. apiocheles. This kind of variability was also described in another endemic species of *Caridina* from the SWIO, *C. similis* (see Marquet & Keith, 2008).

*Caridina similis* Bouvier, 1904

Figs 2C, 5E

*Caridina similis* Bouvier, 1904: 135.

*Caridina brevirostris* f. typica Bouvier, 1912b: 916.

*Caridina brevirostris* f. Gardineri Bouvier, 1912b: 916.

*Caridina similis* – Bouvier 1905: 88, fig. 10. – Keith *et al.* 2006: 58. – Marquet & Keith 2008: 980, figs 1–3.

*Caridina brevirostris* f. similis – Bouvier 1925: 228, figs 507–508.

*Caridina brevirostris* f. typica – Bouvier 1925: 228, figs 509–510, 514–523.

*Caridina brevirostris* f. Gardineri – Bouvier 1925: 228, figs 505–506, 511–513.

*Caridina brevirostris* – Bouvier 1904: 136; 1912b: 915; 1913a: 447, pls. 27–29. — Costa 1980: 673–700.

*Caridina singhalensis* – Borradaile 1907: 67.

**Type locality**

Mahé, Seychelles.

**Material examined**

SEYCHELLES – Mahé • 8 ♀♀ ovig. (cl 4.7–6.3 mm), 1 ♀ (cl 5.9 mm), 1 ♂ (cl 4.1 mm); mare aux cochons; 4°39.029' S, 55°24.152' E; 17 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-128; MNHN • 3 juv. (cl 2.4–2.9 mm); Baume Ellie; 14 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-129; MNHN. – Silhouette • 1 ♀ ovig. (cl 4.4 mm); Anse Mandon; 4°28'1" S, 55°13'30" E; 28 Nov. 2012; P. Keith leg.; DNA voucher: CA1854; MNHN-2025-130; MNHN • 1 ♂ (cl 3.9 mm); same data as for preceding; DNA voucher: CA1855; MNHN-2025-131; MNHN • 1 ♀ ovig. (cl 5.5 mm); same data as for preceding; DNA voucher: CA1856; MNHN-2025-132; MNHN • 1 ♀ ovig. (cl 5.6 mm); same data as for preceding; DNA voucher: CA1857; MNHN-2025-133; MNHN.

See also “other material” in Marquet & Keith (2008).
**Distribution**

Endemic to the Seychelles (La Digue, Mahé, Praslin, Silhouette).

**Habitat**

Freshwater in permanent rivers.

**Remarks**

As for the previous species, *C. similis* shows a very variable form of the rostrum indicating numerous forms like *Caridina brevirostris f. similis*, *Caridina brevirostris f. typica*, *Caridina brevirostris f. Gardineri* now all junior synonyms of *C. similis* (see Marquet & Keith 2008).

*Caridina typus* species group

**Diagnosis**

Robust morphology with a straight rostrum, armed or not on the dorsal margin, without apical teeth, antennal spine fused with inferior orbital angle, antennular peduncle equal or more than half of carapace in length, pterygostomian margin blunt to rather narrowly rounded, stout walking legs, carpus of first pereiopod excavated, short sixth abdominal somite (less than half of carapace length), high pre-anal carina with no spine or a small one, a great number of spinules on the uropodal diaeresis (> 15), plumose terminal setae on the telson subequal to lateral ones or slightly longer, and a long subrectangular endopod of the first male pleopod with a short appendix on the subdistal outer margin.

*Caridina typus* H. Milne Edwards, 1837

Figs 2G, 5F

*Caridina typus* H. Milne Edwards, 1837 (in H. Milne Edwards, 1834–1840): 363, pl. 25bis figs 4–5.

*Caridina exilirostris* Stimpson, 1860: 29.

*Caridina typus f. typica* Bouvier, 1925: 250, figs 272–295.

*Caridina typus f. caledonica* Bouvier, 1925: 253, figs 296–297.

*Caridina typus* – Miers 1879: 492. — Richters 1880: 162, pl. 17 fig. 23. — Pfeffer 1889: 35. — De Man 1892: 367, pl. 21 fig. 22; 1893: 300. — Sharp 1893: 111. — Ortmann 1894a: 8; 1894b: 403. — Weber 1897: 167 (part). — Hilgendorf 1898: 34. — Nobili 1899: 232. — Doflein 1900: 127. — Roux 1904: 552. — Bouvier 1904: 133. — Lenz 1905: 385; 1910: 570. — Borradaile 1907: 67. — Miyake 1938: 111, fig. 1. — Edmondson 1935: 10, fig. 3g–l. — Kubo 1938: 83, fig. 13; 1941: 307. — Kamita 1959: 21; 1961: 45, figs 15–18; 1963: 5, fig. 5; 1967: 3, fig. 4, pl. Ib. — Johnson 1960: 180; 1963: 27; 1967: 418; 1969: 110. — Holthuis 1965: 9; 1969: 93; 1978: 30; 1986: 590. — Ooishi 1970: 87. — Tiwari & Pillai 1971: 186. — Suzuki 1972: 10, figs 8–10; 2001: 138, fig. 2. — Fujino 1972: 7, fig. 8. — Kensley 1972: 36 (key), fig. 16c–d; 1981: 4. — Shokita 1975: 119; 1979: 204, pl. 1e; 1981: 15; 2002: 164; 2003: 249, figs 18g, 19c, 20l. — Shokita & Nishijima 1977: 187. — Costa 1980: 693; 1984: 205. — Hart 1983: 67. — de Silva 1983: 209. — Benzie & de Silva 1984: 623. — Gurney 1984: 568 (key). — Ng 1985: 161; 1995: 189, fig. 7. — Maciolek & Ford 1987: 628. — Coetzee 1988: 62. — Hayashi 1989: 310, fig. 168. — Bright 1989: 34. — Choy 1991: 356. — Hung *et al.* 1993: 486, figs 1c, 4a–f. — Suzuki *et al.* 1993: 56. — Suzuki & Sato 1994: 60. — Hamano & Honke 1997: 162. — Chace 1997: 21. — Shy & Yu 1998: 61. — Yeo *et al.* 1999: 225. — Ebenezer & Richard 1999: 427. — Keith *et al.* 1999: 46; 2006: 62; 2010: 46. — Ideguchi *et al.* 2000: 5. — Short 2000: 61. 2009: 36 (key), fig. 22. — Buden *et al.* 2001: 261. — Leberer & Nelson 2001: 393. — Day *et al.* 2001: 96 (key), fig. 6.3. — Keith 2002: 100. — Choy & Marquet 2002: 219. — Marquet *et al.* 2002: 222; 2003: 68. — Shokita *et al.* 2002: 76; 2003a: 101; 2003b: 117. — Davie 2002: 226.
— Leberer & Cai 2003: 354. — Cai & Anker 2004: 236. — Liang 2004: 139, fig. 65. — Wowor *et al.* 2004: 341, fig. 5f. — Bossuyt *et al.* 2004: 480, fig. 2e. — Nakahara *et al.* 2005: 305. — Cai *et al.* 2006: 412, figs 13–15; 2007: 279; 2009: 66. — Cai & Shokita 2006a: 2134. — Page *et al.* 2007: 647, figs 2–3. — Fujita 2007: 93. — Ragunathan & Valarmathi 2007: 94 (key), 96. — Karge & Klotz 2007: 92. — Nakahara *et al.* 2007: 387, figs 1c, 2a–e. — Cai & Ng 2009: 1094. — Benstead *et al.* 2009: 459. — Cai & Husana 2009: 62. — Richard & Clark 2010: 639. — Soomro *et al.* 2010: 721; 2011: 41; 2016: 569; 2020: 1333. — Saito *et al.* 2012: 27. — Cook *et al.* 2012: 289, fig. 3. — Bouchard *et al.* 2013: 9. — Emmerson 2016: 162, 173. — Fujita *et al.* 2016: 105. — Bernardes *et al.* 2017: 1 (part, ARC clade). — de Mazancourt *et al.* 2017: 226, fig. 4; 2019b: 167, figs 2–5; 2020: 34, figs 2d, 10. — Chow *et al.* 2018: fig. 1. — Choy *et al.* 2019: 115. — Short *et al.* 2019: 4, 20 (key), figs 8, 10j. — Wood *et al.* 2019: 200. — Jugovic *et al.* 2019: fig. 1. — Rahayu & Annawaty, 2019: 64. — Klotz *et al.* 2019: 164, fig. 18. — Hernawati *et al.* 2020: fig. 3. — Imai *et al.* 2020: 63, fig. 3a. — Vijayamma *et al.* 2021: 404 (key).

*Caridina typa* – Bouvier 1905: 88; 1913a: 465; 1914: 699; 1925: 249, figs 271–297. — Roux 1926a: 246; 1926b: 201. — Riek 1953: 117. — Choy & Horwitz 1995: 52 (key) (incorrect spelling).

*Caridina typus* var. *acuminata* – Bouvier 1925: 126, fig. 271 (part).

*Caridina typa acuminata* – Roux 1934: 222.

*Caridina acuminata* – Shokita 1979: 205. — Hayashi 1989: 311, fig. 169b–c, 170f.

? *Caridina aff. typus* – Ng 1994: 78.

Not *Caridina typus* – Spence Bate 1888: 704, pl. 119 fig. 3 (= *Caridina Spencebatei* De Man, 1892: 371 = *Atya scabra*).

Not *Caridina typus* – Weber 1897: 167 (part) (= *Caridina africana*).

Not *Caridina typus* – de Silva 1982: 135, fig. 4a–h (= *Caridina villadolidi*).

Not *Caridina typus* – von Rintelen *et al.* 2007: 1037, fig. 2; 2008: 2248, fig. 4; 2012: figs 2–5.

Not *Caridina typus* – Bernardes *et al.* 2017: 1 (part, SUL clade) (= *Caridina jeani*).

Not *Caridina typus* – Bernardes *et al.* 2017: 1 (part, TAL clade) (= *Caridina zhujiangensis*).

## Type locality

Unknown, probably Mauritius according to Bouvier (1925).

## Material examined

AUSTRALIA • 2 specs from GenBank: DQ478562 and DQ478563 (Page *et al.* 2007).

MADAGASCAR • 1 ♀ (cl 4.5 mm); Antsatoko River; 13°36.660' S, 50°00.341' E; 29 m a.s.l.; 6 Jul. 2008; DIAMSOI leg.; DNA voucher: CA1038; MNHN-IU-2018-2829; MNHN • 1 ♀ ovig. (cl 6.9 mm); same data as for preceding; DNA voucher: CA1040; MNHN-IU-2018-2830; MNHN • 1 ♀ ovig. (cl 7.1 mm); same data as for preceding; DNA voucher: CA1039; MNHN-IU-2018-2831; MNHN.

MAYOTTE • 1 ♀ (cl 6.1 mm); M'Tsangachei River; 12°53.066' S, 45°07.899' E; 30 Mar. 2024; P. Valade leg.; DNA voucher: CA3027; MNHN-IU-2025-134; MNHN.

MAURITIUS • 3 specs from GenBank: KY069380, KY069381 and KY069382 (Bernardes *et al.* 2017).

SEYCHELLES • 1 ♀ (cl 9.2 mm); Praslin Island, Nouvelle Découverte River; 4°19.200' S, 55°42.333' E; 16 m a.s.l.; 8 Oct. 2003; P. Bosc, H. Grondin, P. Keith and P. Valade leg.; MNHN-IU-2014-577; MNHN.

SOUTH AFRICA • 1 ♀ ovig. (cl 10.1 mm); Umtata River; 31°55.511' S, 29°08.199' E; 2 m a.s.l.; 13 Feb. 2018; L. Maliwa, M. Mlambo, G. Marquet and P. Tiberghien leg.; DNA voucher: CA2089; MNHN-
2018-2843; MNHN • 1 ♂ (cl 5.7 mm); Mpenjati River; 30°56.645' S, 30°13.660' E; 93 m a.s.l.; 15 Feb. 2018; P. Kubheka, N. McClurg, T. McClurg, L. Maliwa, M. Mlambo, G. Marquet and P. Tiberghien leg.; DNA voucher: CA2090; MNHN-IU-2018-2844; MNHN.

LA RÉUNION • 1 juv. (cl 2.4 mm); Rivière du Mât; 20°58.801' S, 55°41.989' E; 19 May 2023; P. Valade leg.; DNA voucher: CA3054; MNHN-IU-2025-135; MNHN • 1 ♂ (cl 4.6 mm); same data as for preceding; DNA voucher: CA3058; MNHN-IU-2025-136; MNHN • 1 juv. (cl 2.4 mm); same data as for preceding; DNA voucher: CA3059; MNHN-IU-2025-137; MNHN • 1 juv. (cl 1.9 mm); Rivière des Marsouins; 21°01. 970' S, 55°42.943' E; 6 Aug. 2023; P. Valade leg.; DNA voucher: CA3060; MNHN-IU-2025-138; MNHN • 1 ♂ (cl 4.5 mm); same data as for preceding; DNA voucher: CA3063; MNHN-IU-2025-139; MNHN • 1 spec. from GenBank: KY069434 (Bernardes *et al.* 2017).

**Distribution**

Widespread from South Africa to Fiji and Australia to Japan and India.

**Habitat**

Freshwater in rivers.

**Remarks**

*Caridina typus* revealed to be a species complex, all characterized by the lack of teeth on the dorsal margin of the rostrum. Three clades were recognized by Bernardes *et al.* (2017), named SUL, TAL and ARC. The latter one corresponds to the true *C. typus*, while SUL is *C. jeani* and TAL is *C. zhujiangensis* (publication in prep.).

*Caridina serratirostris* species group

**Diagnosis**

Moderately robust morphology with a moderately short and straight rostrum (reaching end of antennular peduncle), armed with many dorsal teeth, at least 6 of them on the carapace, without apical teeth, the antennal spine ventral to the inferior orbital angle, a long antennular peduncle (about half the carapace length) with an usually long stylocerite (reaching end of first segment of antennular peduncle), pterygostomian margin rounded, segments of walking legs very slender, sixth abdominal somite about half of carapace length, a small pre-anal carina sometimes bearing an acute spine, a great number of spinules on the uropodal diaeresis (> 15), numerous, long and plumose terminal setae on the telson and a rounded endopod of the first male pleopod, without appendix interna.

*Caridina henriettae* sp. nov.

urn:lsid:zoobank.org:act:F229A5EF-5D4B-412D-B6F8-285C1B0B1C27

Figs 2F, 3–4, 5D, G–H

*Caridina serratirostris* – Holthuis 1965: 25, fig. 8. — Costa 1980: 691. — Gurney 1984: 568 (key). — Keith *et al.* 1999: 44; 2006: 56. — Keith 2002: 100. — Richard & Clark 2010: 310 (part), figs 3, 4a–n, p–q. — Bouchard *et al.* 2013: 9. — Emmerson 2016: 170. — Hoarau 2018: 1.

*Caridina wyckii* — Weber 1897: 168 (part).

**Diagnosis**

Typical morphology of the *C. serratirostris* species group: rostrum armed with many dorsal teeth, at least 6 of them on the carapace, a usually long stylocerite and a rounded endopod of the first male pleopod,
without appendix interna. Well-developed arthrobranch at the base of P1. Telson with a pair of plain long lateral spines and 1 pair of shorter, plain lateral spines. P3 propodus 11.2–12.8 as long as wide.

## Etymology

The new species is dedicated to our friend Elvina Henriette, Research Scientist and Director of the NGO “Gaea Conservation Network Seychelles”, in recognition of her work and passion for the freshwater fauna of Seychelles islands.

## Type material

### Holotype

MAURITIUS • 1 ♂ (cl 2.9 mm); Rivière des Anguilles; 20°28.802' S, 57°33.378' E; 13 Nov. 2002; P. Keith, G. Marquet and ARDA leg.; DNA voucher: CA2304; MNHN-IU-2015-1914; MNHN.

### Paratypes

MAURITIUS • 1 ♀ (cl 4.1 mm); same data as for holotype; DNA voucher: CA2305; MNHN-2025-140; MNHN.

MADAGASCAR • 1 ♀ ovig. (cl 4.9 mm); Fanambana River (Station 37); 14°33.193' S, 50°10.393' E; 11 m a.s.l.; 7 Jul. 2008; DIAMSOI leg.; DNA voucher: CA1178; MNHN-2025-141; MNHN • 1 ♂ (cl 3.2 mm); same collection data as for preceding; DNA voucher: CA1431; MNHN-2025-142; MNHN.

MAURITIUS • 1 ♀ ovig. (cl 4.8 mm); Rivière de la Chaux; 20°24.350' S, 57°42.269' E; 12 Nov. 2002; ARDA leg.; DNA voucher: CA2042; MNHN-IU-2015-1933; MNHN • 1 ♂ (cl 3.0 mm); Beau Vallon; 1913; P. Carié leg.; MNHN-IU-2015-1909; MNHN.

LA RÉUNION • 1 ♀ ovig. (cl 4.4 mm); Rivière des Embouchures; 21°2.093' S, 55°42.899' E; 13 Oct. 2015; P. Valade leg.; DNA voucher: CA2417; MNHN-2025-143; MNHN • 1 ♀ ovig. (cl 3.8 mm); same data as for preceding; DNA voucher: CA2418; MNHN-2025-144; MNHN • 1 ♀ ovig. (cl 3.6 mm); Rivière des Marsouins; 2 May 2022; Ocea Consult leg.; DNA voucher: CA2949; MNHN-2025-145; MNHN • 1 ♀ ovig. (cl 3.6 mm); Rivière des Marsouins; 21°1.970' S, 55°42.949' E; 8 Apr. 2024; P. Valade leg.; DNA voucher: CA3038; MNHN-2025-146; MNHN • 1 juv. (cl 2.4 mm); Rivière des Marsouins; 21°1.970' S, 55°42.949' E; 6 Aug. 2023; P. Valade leg.; DNA voucher: CA3061; MNHN-2025-147; MNHN • 1 juv. (cl 2.3 mm); Rivière des Roches; 21°0.398' S, 55°41.661' E; 13 Nov. 2023; P. Valade leg.; DNA voucher: CA3065; MNHN-2025-148; MNHN.

SEYCHELLES – Mahé • 2 ♀♀ ovig. (cl 4.1–4.7 mm), 1 ♀ (cl 4.0 mm); Bel Ombre; 4°36.966' S, 55°25.063' E; 2 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-149; MNHN • 1 ♀ ovig. (cl 2.7 mm); Grand Anse; 4°40.502' S, 55°26.728' E; 20 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-150; MNHN • 1 ♀ ovig. (cl 2.7 mm); same data as for preceding; DNA voucher: CA3000; MNHN-2025-151; MNHN. – Praslin • 1 ♀ ovig. (cl 4.1 mm); 10 Oct. 2003; no locality data; ARDA leg.; DNA voucher: CA2026; MNHN-IU-2015-1932; MNHN • 1 ♀ ovig. (cl 3.4 mm); Nouvelle Découverte; 4°20.179' S, 55°43.349' E; 19 Feb. 2024; P. Keith and M. Mennesson leg.; MNHN-2025-152; MNHN • 2 ♀♀ ovig. (cl 3.2–3.6 mm); Nouvelle Découverte; 4°20.179' S, 55°43.349' E; 21 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-153; MNHN • 1 ♀ ovig. (cl 3.6 mm); same data as for preceding; DNA voucher: CA2998; MNHN-2025-154; MNHN • 1 ♀ ovig. (cl 3.4 mm); same data as for preceding; DNA voucher: CA2999; MNHN-2025-155; MNHN. – Silhouette • 1 ♀ (cl 3.4 mm); Anse Mandon; 4°28.014' S, 53°13.500' E; 28 Nov. 2012; P. Keith leg.; DNA voucher: CA1491; MNHN-2025-156; MNHN • 1 ♀ ovig. (cl 4.5 mm); same data as for preceding; DNA voucher: CA1492; MNHN-2025-157; MNHN • 4 ♀♀ ovig. (cl 3.3–3.7 mm), 1 ♀ (cl 3.3 mm); St Gore River; 4°30.115' S, 55°13.521' E; 18 Feb. 2020; P. Keith and M. Mennesson leg.; MNHN-2025-158; MNHN.
<figure><img src="image_4.png" /><figcaption>**Fig. 3.** *Caridina henriettae* sp. nov. **A–F, H, O.** ♂, holotype (MNHN-IU-2015-1914). **G, J–K.** ♂, paratype (MNHN-IU-2015-1909). **I, L.** Ovigerous, ♀, paratype (MNHN-IU-2015-1933). **M.** Ovigerous female, paratype (MNHN-IU-2025-143). **N.** Ovigerous female, paratype (MNHN-IU-2025-152). **A.** First pereiopod. **B.** Second pereiopod. **C.** Third pereiopod. **D.** Fifth pereiopod. **E.** Dactylus of third pereiopod. **F.** Dactylus of fifth pereiopod. **G.** Uropodal diaeresis. **H.** Telson. **I.** pre-anal carina. **J.** First male pleopod. **K.** Second male pleopod. **L.** Undeveloped eggs. **M–N.** Rostrum variation. **O.** Cephalothorax.</figcaption></figure>
**Description**

CEPHALOTHORAX. Antennal spine ventral to inferior orbital angle. Pterygostomian margin rounded. Rostrum (Fig. 3M–O): straight, short, 0.6–0.7 of cl, reaching beyond end of antennular peduncle, armed with 19–31 teeth on dorsal margin, 6–10 of them situated on carapace behind orbital margin, ventral margin with 5–7 teeth. Eyes developed, anterior end reaching to 0.59 × length of basal segment of antennular peduncle. Long antennular peduncle, 0.67 (♀) – 0.78 (♂) × as long as carapace; second segment distinctly longer than third segment. Stylocerite reaching to middle of second segment of antennular peduncle.

PEREIPODS. Epipods on first four pereiopods. P1 (Fig. 3A): chela about 2.4–3.3 × as long as wide, movable finger 3.7–5.8 as long as wide, 1.4–1.9 × length of palm; carpus 3.6–4.6 × as long as wide, well developed arthrobranch (Fig. 4). P2 (Fig. 3B) more slender and longer than first pereiopod, with chela 3.7–5.2 × as long as wide: movable finger 5.2–11.0 × as long as wide, 1.4–2.4 × as long as palm; carpus slender, 7.7–9.6 × as long as wide. P3 (Fig. 3C): slender, dactylus (Fig. 3E) 3.2–4.0 × as long as wide (terminal spiniform seta included), with 5–8 spiniform setae on flexor margin including terminal one; propodus 11.2–12.8 × as long as wide, 3.2–4.4 × as long as dactylus. P5 (Fig. 3D): dactylus (Fig. 3F) 3.9–4.4 as long as wide, with 7–13 spiniform setae on flexor margin; propodus 15.8–23.9 × as long as wide, 4.4–50 × as long as dactylus.

ABDOMEN. Third abdominal somite with moderately convex dorsal profile. Sixth abdominal somite 0.49 × as long as carapace, 1.3 × as long as fifth somite, shorter than telson.

<figure><img src="image_5.png" /><figcaption>**Fig. 4.** Dissected branchial chamber of a specimen of *Caridina henriettae* sp. nov., paratype (MNHN-2025-149) showing the developed arthrobranch at the base of the first pereiopod (black arrow).</figcaption></figure>
<figure><img src="image_6.png" /><figcaption>**Fig. 5.** Live coloration of the species of Atyidae from the South West Indian Ocean. A. *Atyoida serrata* Bate, 1888 from La Réunion (credit E. Vigneux). B. *Caridina natalensis* De Man, 1908 from the Seychelles (credit P. Keith). C. *Caridina spathulirostris* Richters, 1880 from Mauritius (credit Hoareau). D. *Caridina henriettae* sp. nov. from the Seychelles (credit P. Keith). E. *Caridina similis* Bouvier, 1904 from the Seychelles (credit Y. Zitte). F. *C. typus* H. Milne Edwards, 1837 from La Réunion (credit E. Vigneux). G–H. *C. henriettae* sp. nov. from La Réunion (credit E. Vigneux).</figcaption></figure>
TELSON (Fig. 3H). 3.0 × as long as wide, with 6 pairs dorsal spines including subterminal spines, 1 pair of plain long lateral spines, 1 pair of shorter plain lateral spines and 6–11 long, stalked, plumose processes present at rounded posterior margin.

MALE PLEOPODS. P11 (Fig. 3J): endopod rounded, 2.0 × as long as wide, reaching 0.35 × as long as endopod, no appendix. P12 (Fig. 3K): appendix masculina reaching 0.74 × as long as endopod; appendix interna reaching 0.82 × as long as appendix masculina.

PRE-ANAL CARINA (Fig. 3I). High, with a spine.

UROPODAL DIAERESIS (Fig. 3G). With 13–15 spinules.

EGGS. Ovigerous females with small eggs (Fig. 3L), size: 0.37–0.46 × 0.24–0.27 mm.

## Distribution

South Africa, Mozambique, Madagascar, Seychelles, Comoros, La Réunion, Mauritius.

## Habitat

Fresh or brackish water in the lower course and estuary of rivers.

## Remarks

This new species belongs to the *Caridina serratirostris* species group based on the presence of a rostrum armed with many dorsal teeth, at least 6 of them on the carapace, with a usually long stylocerite and with a rounded endopod of the first male pleopod, without appendix interna. But it is easy to distinguish this new species from type specimens of congeners by its telson with a pair of plain long lateral spines and 1 pair of shorter, plain lateral spines (vs only a pair of plain lateral spines) and by its P3 propodus shorter 11.2–12.8 × as long as wide (vs 14.2–17.1). Among this group, our new species is closer to *C. serratirostris* than *C. celebensis* due to the presence of a well-developed arthrobranch at the base of the P1 (vs reduced or absent, see Cai & Shokita 2006b)

Richard & Clark (2010) reported this species under the name *C. serratirostris* from Mozambique and South Africa noting the presence of shorter inner lateral spines on the telson which the syntypes of *C. serratirostris* do not possess. Recent sampling trips to the localities where the species was collected in South Africa – the iLovu or the Umbilo rivers near Durban – by two of the authors (VM and GM) failed to retrieve this species, suggesting either that this area is the southern limit of its range and its presence may be occasional, or that the species does not occur there anymore due to the heavy urbanization around these rivers (pers. obs.).
## Identification key for the species of *Caridina* H. Milne Edwards, 1837 from the South-Western Indian Ocean

1. Stylocerite reaching distinctly beyond the base of the second segment of the antennular peduncle .................................................................................................................. *C. henriettae* sp. nov.
– Stylocerite at most attaining the end of the first segment of the antennular peduncle ........................................ 2

2. Rostrum rather constantly long, reaching well beyond scaphocerite .... *C. natalensis* De Man, 1908
– Rostrum short or reaching at most to or slightly beyond the end of the antennular peduncle ............ 3

3. Uropodal diaeresis with 16–23 spinules .................................................................................................................... 4
– Uropodal diaeresis with 8–15 spinules .................................................................................................................... 5

4. Dactylus of P5 with 26–37 spiniform setae; ovigerous female with few and large eggs ....................................................
– Dactylus of P5 with 33–81 spiniform setae; ovigerous female with numerous and small eggs ....................
*C. similis* Bouvier, 1904 (Seychelles)
*C. typus* H. Milne Edwards, 1837

5. Dorsal margin of telson unarmed; dactylus of P5 with 28–30 spiniform setae ....................................................
– Dorsal margin of telson armed; dactylus of P5 with 31–60 spiniform setae ....................................................
*C. mauritii* Bouvier, 1912 (Mauritius)
*C. spathulirostris* Richters, 1880 (Mauritius)

## Discussion

This updated checklist of the species of *Caridina* from the SWIO gives new insights in the biogeography of the region. Some of the species listed have a wide distribution range due to their amphidromous life cycle: *C. typus* is one of the most widely distributed species of the genus being able to cross the Indo-Pacific barrier. *Caridina natalensis* and *C. henriettae* sp. nov. have a smaller range, being restricted to the SWIO, probably because of a shorter planktonic larval phase (de Mazancourt *et al.* 2023). Endemic land-locked species can be found in the Seychelles with *C. similis* and Mauritius with *C. mauritii* and *C. spathulirostris*, without mentioning the large endemic species flocks from Madagascar that were intentionally left aside from the present study due to the lack of data. The absence of such species in La Réunion Island while two co-exist in the neighboring island of Mauritius can be explained by the older age of the latter (about 10 My vs 3 My) which gave enough time for endemic species to evolve. Similarly, the granitic islands of the Seychelles were formed very early from a microcontinent separated from India in the late Cretaceous (66 My), which also explains the presence of endemic species like *C. similis* or the primary freshwater crab genus *Seychellum* (Ng *et al.* 1995).

## Acknowledgments

For the Seychelles, we would like to thank particularly our friends E. Henriette and Wilna Accouche from Gaea who organized very efficiently the two trips in 2020 and 2024 with their team and particularly for their help on the field Gina Cesar, Staniella Henriette, Mersiah Rose, and Terence Vel; thanks to the PS Alain Decomarmond, Department of Environment, Mrs Marie-May Muzungaile and Ashley Dias, Biodiversity Conservation and Management Division, Elvis Nicette, Seychelles Bureau of Standards, Mr Daniel Etongo and Nuette Gordon, University of Seychelles; thanks to the Environment, Energy & Climate Change (MEECC), for agreeing to the transfer of material; thanks also to all people who join us in the field. For the Comoros, we would particularly like to thank Yahaya Ibrahim (CNDRS). For Madagascar, we are grateful to J. Aride from Madagascar National Parks (MNP-ANGAP), to the Manager of Masoala National Parc at Maroantsetra, and to the DIAMSOI team (research financed by the Institut Français de la Biodiversité (IFB)), and particularly to his coordinator E. Feunteun (MNHN). For
La Réunion Island and Mayotte, we thank the Environnement Offices, Ocea Consult (P. Valade), Arda and the Agriculture Offices. For Mauritius, thanks to the CWA, M. Rohit Mungra, M. Dev Aukle & M. Anand Kumar Gopaul, the Ministry of Environment, M. Phosun Kallee and the NEL, M. Fareed & M. Igbal Atchia, the WRU, M. Hari Ram Sharma, M. Farook Mowlabucus, Ms Suzanne Boodhoo & M. R. Teeluck. For South Africa, we are grateful to Patrick Kubheka and Helen James for facilitating collecting permits for us with provincial conservation urgencies in KwaZulu-Natal and Eastern Cape (permit no. CRO69/17CR), respectively. We thank Nick and Tim McClurg, Patrick Kubheka, Lazola Maliwa and Pascal Tiberghien for their help in the field and Albert Chakona (SAIAB) for lending us his electric fishing gear. Thanks to the Service de Systématique Moléculaire of the MNHN. Finally, we wish to thank Werner Klotz for his comments that helped improving the quality of the manuscript.

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Printed versions of all papers are deposited in the libraries of four of the institutes that are members of the EJT consortium: Muséum national d'Histoire naturelle, Paris, France; Meise Botanic Garden, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences, Brussels, Belgium. The other members of the consortium are: Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo Nacional de
Ciencias Naturales-CSIC, Madrid, Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the Czech Republic, Prague, Czech Republic; The Steinhardt Museum of Natural History, Tel Aviv, Israël.