Introduction
Amphipods of family Caprellidae Leach, 1814, also known as ‘skeleton shrimps’ are generally found associated with a variety of substrates such as algae, bryozoans, sponges, and hydroids (McCain, Reference McCain1968; Guerra-García, Reference Guerra-García2004; Momtazi et al., Reference Momtazi, Sari and Darvish2020). Caprellids also exhibit notable ecological adaptability, with distribution and abundance of many species strongly influenced by habitat structure, hydrodynamic conditions and environmental gradients (Woods, Reference Woods2009). These ecological factors often drive morphological variations complicating species-level identification.
The genus Caprella can be identified by the following characters: Pereopod 3 and 4 absent; gills present on pereonite 3 and 4; mandible without palp; male with a pair of abdominal appendages (Arimoto, Reference Arimoto1976). Caprellidae comprises more than 430 species out of which around 185 species belong to genus Caprella Lamarck, 1801 only, making it the most diverse genus among the family (Horton et al., Reference Horton, Lowry, De Broyer, Bellan-Santini, Copilas-Ciocianu, Corbari, Costello, Daneliya, Dauvin, Fišer, Gasca, Grabowski, Guerra-García, Hendrycks, Hughes, Jaume, Jazdzewski, Kim, King, Krapp-Schickel, LeCroy, Lörz, Mamos, Senna, Serejo, Souza-Filho, Tandberg, Thomas, Thurston, Vader, Väinölä, Valls Domedel, Vonk, White and Zeidler2025). From India, 18 species of family Caprellidae have been reported, among these four species belong to genus Caprella: C. danilevskii (Czerniavski, Reference Czerniavski1868); C. madrasana Giles, 1888; C. mendax Mayer, 1903, and C. palkii Giles, 1888 (Thacker et al., Reference Thacker, Patel, Myers, Guerra-García, Zeidler and Trivedi2023a).
Although C. danilevskii was originally described from Black Sea, it has been reported widely beyond its type locality, resulting in taxonomic confusion with morphologically similar species. Caprella moradi was subsequently described from the Gulf of Oman based on consistent morphological differences from C. danilevskii. In this study, newly collected material from the same locality reported by Gaikwad and Sautya (Reference Gaikwad and Sautya2022) was examined and comparison with their published figures indicates that previously reported specimens from Gujarat, India correspond to C. moradi. Therefore, we consider the earlier Indian record of C. danilevskii to be synonymous with the C. moradi.
Gujarat State, situated in the western part of India, possesses the longest coastline in the country, stretching approximately 2340 km (Patel et al., Reference Patel, Rabari, Patel, K and Trivedi2025). This extensive coastline encompasses diverse marine habitats, including sandy shores, rocky shores, mudflats, coral reefs, mangroves, and estuaries, which together sustain a rich diversity of crustacean fauna (Gosavi et al., Reference Gosavi, Trivedi, Trivedi and Vachhrajani2017). While the diversity of crustacean groups such as brachyurans, anomurans, and stomatopods along the coast of Gujarat has been thoroughly investigated (Trivedi et al., Reference Trivedi, Soni and Vachhrajani2015, Reference Trivedi, Ahyong, Vachhrajani and Kumar2020; Trivedi and Vachhrajani, Reference Trivedi and Vachhrajani2017; Gosavi et al., Reference Gosavi, Purohit, Mitra, Patel, Vachhrajani and Trivedi2021; Patel et al., Reference Patel, Padate, Osawa, Tiwari, Vachhrajani and Trivedi2022), studies focusing on amphipod fauna are comparatively scarce (Myers et al., Reference Myers, Trivedi, Gosavi and Vachhrajani2017, Reference Myers, Trivedi, Gosavi and Vachhrajani2018; Gaikwad and Sautya, Reference Gaikwad and Sautya2022; Thacker et al., Reference Thacker, Myers and Trivedi2023b, Reference Thacker, Myers and Trivedi2023c, Reference Thacker, Myers and Trivedi2024a, Reference Thacker, Myers, Trivedi and Mitra2024b, Reference Thacker, Myers and Trivedi2024c, Reference Thacker, Myers and Trivedi2025a, Reference Thacker, Myers and Trivedi2025b). Approximately 265 species of amphipods have been documented in India, with only 26 species reported from Gujarat (Thacker et al., Reference Thacker, Patel, Myers, Guerra-García, Zeidler and Trivedi2023a). In this study, C. moradi is reported for the first time from Gujarat, India, adding a new addition to the amphipod checklist for both Gujarat and India.
Materials and methods
Specimens examined for the present study were collected from Diu (20°42’06.8”N 70°55’01.2”E) and Veraval (20°54’35”N 70°21’09”E), Gujarat. Sampling was carried out in the intertidal zone, where specimens were found attached to the algal substrates dominated by Chaetomorpha spp. and Cystoseira spp. occurring over rocky substratum. During sampling, seawater temperature was around 26–28 °C and salinity was around 34–35 PSU. A total of 10 samples were collected, with sizes ranging from 5.0 to 10.0 mm. The collection process involved the careful use of forceps to pluck the specimens directly from the algal substrate. Once collected, the specimens were transferred into the specimen vials containing 5% formaldehyde solution. Then the vials were subsequently labelled with details of the collection site and date. Later in the laboratory the specimens were transferred to 70% ethanol. Dissection of different body parts was carried out using a stereomicroscope (Metlab PST-901) for species-level identification. After dissection photographs were captured using camera attached to stereomicroscope (Trinocular Stereo Zoom Microscope, magnification of 7×–45×) and detailed illustrations were prepared by tracing the photographs in the Inkscape (an open-source vector graphics editor) software. Type materials are deposited in LFSC.ZRC, Department of Life-sciences, Hemchandracharya North Gujarat University, Patan, with reference number (LFSC.ZRC-289).
Results
Order Amphipoda Latreille, 1816
Suborder Senticaudata Lowry & Myers, 2013
Family Caprellidae Leach, 1814
Genus Caprella Lamarck, 1801
Caprella moradi Momtazi, Sari & Darvish, 2020
(Figs. 2–4)
Caprella moradi: Momtazi et al., 2020: 562–571, figs. 1–6.
Caprella danilevskii: Gaikwad & Sautya, 2022: 33–39, figs. 1–6.
Material examined. 8 males, 5.0–10.0 mm (LFSC.ZRC-289), Diu (20°42’06.8”N 70°55’01.2”E), Gujarat, India, 20 January, 2023, coll. D.R. Thacker; 2 males, 6.0 & 7.0 mm (LFSC.ZRC-289), Veraval, 9 February, 2023, Coll. D.R. Thacker.
Diagnosis. Body surface covered with four rows of small setae (Figure 3H). Male gnathopod 2 propodus anterior margin curved (Figure 4B). Gills 3 and 4 unequal in size (Figure 4C & D).
Map showing the known geographical distribution of Caprella moradi (Momtazi et al., Reference Momtazi, Sari and Darvish2020), including the type locality (Momtazi et al., Reference Momtazi, Sari and Darvish2020) and records obtained in the present study.

Caprella moradi (Momtazi et al., Reference Momtazi, Sari and Darvish2020), male, 7.5 mm (LFSC.ZRC-289). Scale bar = 1.0 mm.

Caprella moradi (Momtazi et al., Reference Momtazi, Sari and Darvish2020), male, 8.2 mm (LFSC.ZRC-289). (A) Antenna 1; (B) Antenna 2; (C) Maxilla 1; (D) Maxilla 1; (E) Maxilliped; (F) Labrum; (G) Labium; (H) four rows of small setae (only 3 visible here from the side view); (I) abdominal appendages. Scale bars = 0.25 mm.

Caprella moradi (Momtazi et al., Reference Momtazi, Sari and Darvish2020), male, 8.2 mm (LFSC.ZRC-289). (A) Gnathopod 1; (B) Gnathopod 2; (C) Gill 3; (D) Gill 4; (E) Pereopod 5; (F) Pereopod 6; (G) Pereopod 7. Scale bars = 0.25 mm.

Description. Based on male 8.2 mm.
Head. Antenna 1 slender, length ratio of peduncular articles 1–3 1:1.63:1; flagellum with 10–11 articles; 0.35× as long as peduncle (Figure 3A). Antenna 2 stout, length ratio of peduncular articles 3–5 1:1.23:0.7; flagellum uni-articulate, articles 3 and 4 with long setae (Figure 3B). Labrum notched distally, 1.12× as long as broad, small setae on distal margin (Figure 3G). Maxilla 1 outer plate with 7 apical robust setae; palp article 2 with 4 triangular projections, 5 apical robust setae and 8–10 simple facial setae (Figure 3C). Maxilla 2 inner plate apically setose, oblique setal row absent; outer plate apically setose (Figure 3D). Maxilliped inner plate with several apical plumose setae; outer plate with 1 apical and 2 marginal robust setae, several marginal simple setae and few facial setae; palp article 4 longest; with an apical nail (Figure 3E).
Pereon. Gnathopod 1 basis longer than wide, proximally wider and narrower towards distal end, anterior margin bare, posterior with several small setae; posterior margin with a single simple seta; ischium as long as broad; merus 1.25× as long as ischium; carpus subtriangular; propodus sub-pyriform, posterior margin serrated, postero-proximal corner developed; dactylus subequal to propodus, inner margin serrated (Figure 4A).
Gnathopod 2 basis slender, bare, 0.8× as long as propodus, both margins bare; merus 1.25× as long as ischium; carpus small, triangular; propodus 4.18× as long as broad, broader distally; posterior margin convex, anterior margin with a triangular process near distal end; dactylus curved, small, reaching 0.33× length of the propodus (Figure 4B).
Pereonite 3 and 4 with gills. Gill 3 suboval, 1.75× as long as gill 4. Gill 4 smaller, suboval. Both gills with small setae surrounding the margins. Pereopod 3 and 4 absent (Figure 4C & D).
Pereopod 5 well developed with six article; basis as long as broad, posterodistal corner produced; ischium small with dorsal carina; merus as long as broad; distal margin produced medially; propodus subrectangular, longer than broad, without robust seta; dactylus curved, inner margin serrated, both margins with few small setae (Figure 4E). Pereopod 6 similar to pereopod 7 but slightly longer (Figure 4F). Pereopod 7 basis longer than broad; merus longer than broad with posterodistal corner produced; dactylus curved, inner margin serrated, both margins with few small setae (Figure 4G).
Pleon. Abdomen with a pair of hooked appendages, a pair of lateral lobes, and one dorsal rounded lobe. Appendages with few setae and 2–3 rows of robust setae on the ventral side. Lateral lobes thin, longer than the medial lobe (Figure 3I).
Remarks. Specimen examined in the present study show agreement with the description and illustrations of C. moradi given by Momtazi et al (Reference Momtazi, Sari and Darvish2020). However, it exhibits certain variations from the type specimen of C. moradi in following characters: abdominal lateral lobes are longer than the dorsal lobe (Figure 3I) whereas, in specimen from Iran (Momtazi et al., Reference Momtazi, Sari and Darvish2020) have lateral lobes shorter than the dorsal (fig. 3AbV); specimen examined here have rounded dorsal lobe (Figure 3I) whereas, in the specimen from Iran (Momtazi et al., Reference Momtazi, Sari and Darvish2020) the dorsal lobe is notched (fig. 3AbV). These variations could be due to the size difference in the examined specimens, as the specimen examined here is larger in size (8.2 mm) than the Iran specimen (6.59 mm) (Momtazi et al., Reference Momtazi, Sari and Darvish2020).
Caprella moradi shows similarity with its congener C. danilevskii in characters such as: absence of grasping spine in pereonite 5–7, short dactylus of male gnathopod 2 and a pair of hooked abdominal appendages. However, C. moradi can be differentiated from C. danilevskii by the presence of four rows of small setae on body surface (versus presence of small setae but without regular placement in C. danilevskii) and curved anterior surface of male gnathopod 2 propodus (versus straight anterior surface of male gnathopod 2 propodus in C. danilevskii). Although C. danilevskii was originally described from the Black Sea (Czerniavski, Reference Czerniavski1868), it is widely distributed and a well-documented member of the genus Caprella in East Asia, where it has been extensively studied taxonomically and ecologically (Vassilenko, Reference Vassilenko1974; Arimoto, Reference Arimoto1976; Takeuchi et al., Reference Takeuchi, Kuwabara, Hirano and Yamakawa1987, Reference Takeuchi, Matsumasa and Kikuchi2003; Lee and Hong, Reference Lee and Hong2011). The species has also been reported from the Atlantic Ocean (McCain, Reference McCain1968), the Mediterranean Sea (Krapp-Schickel, Reference Krapp-Schickel and Ruffo1993) and the Indian Ocean (Mayer, Reference Mayer1890), with intraspecific morphological variation documented among populations from these regions. Caprella moradi was subsequently described from the Gulf of Oman, Iran, based on consistent morphological differences from C. danilevskii (Momtazi et al., Reference Momtazi, Sari and Darvish2020).
Gaikwad and Sautya (Reference Gaikwad and Sautya2022) earlier documented C. danilevskii from Veraval, Gujarat, India. The identification appears questionable, as the anterior margin of the gnathopod 2 propodus in C. danilevskii is straight, whereas the illustrations by Gaikwad and Sautya (Reference Gaikwad and Sautya2022) depict a curved anterior margin (Figure 2F) resembling that of C. moradi. Additionally, the dorsal lobe of the abdominal appendages is notched in the illustrations given by Gaikwad and Sautya (Reference Gaikwad and Sautya2022) (Figure 3K), which also resembles C. moradi (fig. 3AbV). These similarities suggest that the specimen documented previously from Gujarat by Gaikwad and Sautya (Reference Gaikwad and Sautya2022) may, in fact, be C. moradi.
Ecology. Caprellids are marine benthic crustaceans found inhabiting various substrates such as algae, bryozoans, sponges and hydroids (McCain, Reference McCain1968; Guerra-García, Reference Guerra-García2004; Momtazi et al., Reference Momtazi, Sari and Darvish2020) and are also the part of fouling communities (Thiel et al., Reference Thiel, Guerra-García, Lancellotti and Vásquez2003; Ros and Guerra-García, Reference Ros and Guerra-García2012). Caprellid are an important food source of many fish species (Caine, Reference Caine1987, Reference Caine1989, Reference Caine1991). These small crustaceans form an important link between the primary producers such as algae and higher trophic taxa. In the present study, specimens were found attached to algal substrates dominated by Chaetomorpha spp. and Cystoseira spp., occurring over rocky substratum where seawater temperature ranged between 26 and 28 °C and salinity was around 33–34 PSU. Caprella moradi previously was only reported from the Gulf of Oman, Iran, and now it is first time reported from Gujarat State, India, in this study. We hypothesis that the reason behind the range extension of the species from Iran to India could be due to natural or anthropogenic sources such as any floating object or vessels.
Distribution. Djod village, Gulf of Oman, Iran (Momtazi et al., Reference Momtazi, Sari and Darvish2020) Diu and Veraval, Gujarat (this study), India. In this study, this species is first time reported from India (Figure 1).
Acknowledgements
The first author extends heartfelt thanks to IDEA WILD, a non-profit organization for providing small equipment and supplies in support of this biodiversity conservation work.
Author contributions
D.T. prepared the morphological description and illustrations. J.T. reviewed and gave important comments on manuscript. Both authors read and approved the final manuscript.
Funding
This research received no specific grant from any funding agency, commercial or not-for-profit sectors.
Competing interests
The authors declare no conflict of interests.
Data availability
Data will be made available on request.