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18 December 2025

28 Pages

Annotated Checklist and Biodiversity of Sea Cucumbers (Holothuroidea) in Indian Waters

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,
and
1
Graduate School of Integrated Science and Technology, Nagasaki University, 1–14 Bunkyomachi, Nagasaki 852-8521, Japan
2
Department of Ocean Studies and Marine Biology, Pondicherry University, Off Campus, Port Blair 744112, India
3
CSIR National Institute of Oceanography, Regional Centre, Mumbai 400053, India
4
Organization for Marine Science and Technology, Nagasaki University, 1–14 Bunkyomachi, Nagasaki 852-8521, Japan
Oceans2025, 6(4), 85;https://doi.org/10.3390/oceans6040085
This article belongs to the Topic Conservation and Management of Marine Ecosystems
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Abstract

Holothuroids play a vital role in nutrient cycling and bioturbation to enhance the marine ecosystem. They enhance the biodiversity for various symbiotic marine organisms by providing essential shelter and spawning grounds. This review focuses on the Class Holothuroidea (Phylum Echinodermata) in Indian waters, encompassing a total of 187 species organized into 7 orders and 21 families. Notably, the order Holothuriida represents the largest proportion of species, accounting for 27%. These species are well-distributed across India, with the Andaman and Nicobar Islands exhibiting the highest level of species richness (107 species), followed by the East Coast (102 species), Lakshadweep (39 species), and the West Coast (34 species). Species diversity was assessed using the Shannon–Weiner diversity index. Results indicate that the Andaman and Nicobar Islands (H’ = 2.23) and the West Coast (H’ = 2.14) demonstrate the highest levels of diversity. This review provides a comprehensive and precise inventory of all species of Holothuroidea reported in Indian waters, which is provided to facilitate understanding of the reported species, their systematics, and distribution. In addition, a significant insight for both conservation and management of sea cucumbers in India has also been provided.

1. Introduction

Sea cucumbers (Echinodermata, Holothuroidea), comprising approximately 1800 species, inhabit oceans worldwide, thriving in environments ranging from shallow intertidal zones to the ocean’s depths [1,2,3]. A large-scale investigation into echinoderm taxonomy began in the 19th century, utilizing samples collected from expeditions such as Valdivia, Siboga, Challenger, and Albatross [4]. Koehler & Vaney made notable collections of Holothuroidea in the years 1905 [5] and 1908 [6]. Several noteworthy contributions to the current understanding of Indian holothuroid biodiversity have been made [7,8,9,10,11]. These contributions include the documentation of distribution patterns, the identification of taxonomic errors, and the description of new species.
Most commercially exploited holothuroids are typically found in shallow waters, generally at depths of up to 20 m [12]. However, some species, such as Holothuria spinifera, can be located in slightly deeper environments, while Bohadschia marmorata inhabits lagoons. Actinopyga mauritiana is predominantly found in reef areas, where it attaches its tube feet to the substrate [13]. Conversely, the genus Actinopyga species mostly occur in shallow water [14]. Actinopyga echinites occurs mainly on shallow reef flats, often in the presence of live corals [12]. However, it can also be found in intertidal regions. Thelenota ananas resides on reef slopes, coral patches, and hard bottoms at around 30 m depth [12,13].
On the other hand, holothuroids are not only extensively used in the food industry, where they are traditionally processed into beche-de-mer or trepang [15], but also utilized in several cosmetics, nutraceutical, and pharmaceutical industries [16]. Their bodies are rich in various bioactive compounds such as phenols, collagen, amino acids, saponins, and chondroitin sulfate, making them valuable in both the cosmetic and pharmaceutical sectors [16,17,18,19,20].
The Asian waters exhibit a remarkable level of sea cucumber diversity, with over 125 species documented, including 52 that are considered commercially important [21]. This diversity varies across different countries. In China, 134 species are reported, including 27 that are commercially significant [22], 13 species from Taiping Island [23], 30 species from the Taiwan Province of China [24], 25 species from Vietnam [25], 56 species from Indonesia [26]. 62 species from Peninsular Malaysia [27] and Sabah [28], 71 species from Thailand [28,29], and 14 species from North Male Atoll in the Maldives [30]. The lagoons of Mauritius have a higher abundance of holothuroids, with 76% of them commercially important sea cucumbers, where Holothuria atra was found to be the dominant species [31,32]. Meanwhile, Mahdy et al. [33] reported a high abundance of Holothuria atra along the coast of Egypt. Additionally, in another study from Pakistan [34] reported that the distribution and abundance of holothuroids are particularly high along the coast of Karachi. Similarly, Moazzam [35] identified 20 species of holothuroids in addition from the Pakistan coastline, with Holothuria (Theelothuria) hamata being a newly reported species. Clark and Rowe [36] reported 75 species from the intertidal regions of Sri Lanka; however, the recent study by Dissanayake and Stefansson [37] revealed another 25 species from the Sri Lankan coast.
In India, sea cucumbers are primarily found in the Andaman and Nicobar Islands, the Gulf of Kutch, the Lakshadweep Islands, and the Gulf of Mannar region, with smaller populations in other coastal areas. About 12 species are commercially important for beche-de-mer [38]. Sea cucumber fisheries and fisheries trade were previously restricted to the Gulf of Mannar and Palk Bay, where harvesting has been practiced for a long time, primarily targeting species such as Holothuria scabra and H. spinifera [39,40]. However, overexploitation of sea cucumbers during the 1970s and 1980s led to a significant decline in their populations. In response to this issue, regulations were implemented to protect these species by prohibiting the capture of individuals smaller than 75 mm in size. Unfortunately, this measure proved ineffective, prompting the inclusion of all sea cucumber species under Schedule I of the Wildlife Protection Act of 1972 in 2001 [41,42].
This review aims to identify any previously undocumented species in Indian reports, thereby enriching our understanding of regional biodiversity. The review also addresses nomenclatural criteria, evaluates the extent of taxonomic inflation, and updates the checklist of class Holothuroidea from India. It encompasses information on the ecology, biodiversity, in particular distribution, and abundance, making it a valuable resource for researchers and conservationists engaged in marine biodiversity studies.

2. Materials and Methods

Records detailing the distribution and abundance of holothuroids in India were collected from a variety of sources, including websites, checklists, testimonials, articles, and other available resources. A thorough literature review was performed using databases such as ScienceDirect, Google Scholar, ResearchGate, and Scopus. The search utilized keywords including “Holothuroidea”, “sea cucumber diversity”, “Echinodermata”, “Indian Ocean”, “Gulf of Mannar”, “Andaman and Nicobar Islands”, “Lakshadweep”, “Gulf of Kutch”, and “species richness”. Only studies that identified species at the species level and specified the exact locations of their collection were included in the review. For each reported species, scientific names were verified using the WoRMS database. All corrected names were subsequently updated, resulting in the creation of a new checklist. The author meticulously reviewed scientific publications from 1887 to 2024 to ensure the accuracy and currency of the checklist. The checklist reflects significant updates on nomen dubium, unaccepted names, synonyms, and other critical taxonomic terms following the WoRMS database [43]. The data on abundance were irregularly listed across the sources; thus, we constructed presence/absence matrices for each distributed region in their multivariate analyses. These analyses were conducted using PRIMER v7 software. Non-metric multidimensional scaling (nMDS) was utilized to visualize compositional similarity among regions. The nMDS ordination yielded a stress value of zero, indicating a perfect representation of the similarity matrix in two-dimensional space and confirming that the assumptions of nMDS were fully satisfied. Data handling, tabulation, and preliminary calculations were carried out using Microsoft Excel. To interpret the diversity, dominance, and similarity, we organized the data based on different regions, such as the West Coast, East Coast, Andaman and Nicobar Islands, and Lakshadweep (Figure 1).
Figure 1. Map showing the Location of the West Coast, the East Coast, the Andaman and Nicobar Islands, and the Lakshadweep in India. Stars indicate the coastal cities of India.

3. Results

Diversity

The prevalence of sea cucumbers along the coastlines of the Indian Exclusive Economic Zone (EEZ) is attributed to the diverse habitats that support their thriving populations. An updated checklist reveals that a total of 187 species has been identified in Indian waters through previous research, encompassing 7 orders and 21 families. The Andaman and Nicobar Islands exhibit the highest number of species, totaling 108, attributed to their remarkable biodiversity. The East Coast ranks as the second richest marine biodiversity region, harboring approximately 101 species. A significant number of these species have been recorded in the Gulf of Mannar area. After this, Lakshadweep has been documented to host a total of 42 recorded species. The coast with the least biodiversity was the west coast, with a total of 34 species recorded. Among these, the highest number was found along the coast of Kerala. Additionally, 11 species had been reported from India; however, the exact locality of their presence remains unknown. The updated checklist of sea cucumbers in the Indian waters is given below (Table 1). In addition, we have provided several sea cucumber species photographed from the South Coast of Andaman, India (Figure 2 and Figure 3).
Table 1. Updated checklist of the Class Holothuroidea from the Indian waters.
Figure 2. Sea Cucumber along the south coast of the Andaman: (a) Holothuria atra Jaeger,1833, (b) Stichopus chloronotus Brandt, 1835, (c) Actinopyga mauritiana (Quoy & Gaimard, 1834), (d,e) Holothuria (Lessonothuria) pardalis Selenka,1867, (f) Labidodemas semperianum Selenka, 1867, (g) Holothuria leucospilota (Brandt, 1835), (h) Holothuria edulis Lesson, 1830, (i) Holothuria (Thymiosycia) impatiens (Forsskål, 1775).
Figure 3. (a) Stichopus vastus Sluiter,1887, (b) Holothuria (Stauropora) pervicax Selenka, 1867, (c) Stichopus herrmanni Semper, 1868, (d) Bohadschia atra Massin, Rasolofonirina, Conand & Samyn,1999, (e) Stichopus horrens Selenka,1867, (f) Holothuria flavomaculata Semper, 1868, (g) Synapta maculata (Chamisso & Eysenhardt, 1821), (h) Opheodesoma grisea Semper, 1867.
Among these recorded sea cucumbers, the order Holothuriida represents the highest proportion, comprising 27% of the species, followed by Dendrochirotida at 23%. The least reported species belong to the order Persiculida, which constitutes only 3% of the total (Figure 4). Notably, five species, namely Holothuria atra, Holothuria (Lessonothuria) pardalis, Holothuria leucospilota, Holothuria cinerascens, and Stichopus herrmanni, were observed across all selected coastal zones in this review.
Figure 4. Order-wise percentage contribution of Class Holothuroidea.
The dominance of 176 species, excluding the 11 species with uncertain locations, is provided in Figure 5, which encompasses the West Coast, East Coast, Andaman & Nicobar Islands, and Lakshadweep. The station with lower abundance and higher diversity is represented by the lower line. In the k- k-Dominance curve analysis, the Andaman and Nicobar Islands represented a high degree of species dominance, followed closely by the East Coast of India. At the same time, the West Coast exhibited the lowest levels of species dominance.
Figure 5. Species-wise k-Dominance curve using data of the published studies from the West Coast, East Coast, the Andaman & Nicobar Islands, and Lakshadweep.
The Shannon–Weiner diversity index (H’) provides a comprehensive measure of biodiversity by considering both the number of species present and their relative abundance, allowing for a more accurate assessment of overall diversity across research sites. The highest diversity is observed in the Andaman and Nicobar Islands (H’ = 2.23) and the West Coast (H’ = 2.14). The East Coast shows modest variety (H’ = 2.06). In contrast, Lakshadweep Island has relatively low diversity (H’ = 1.39). The Andaman and Nicobar Islands and the West Coast, with their rich biodiversity and favorable environmental conditions, offer conducive habitats for a wide range of species, resulting in higher diversity indices (Table 2).
Table 2. Diversity indices of class Holothuroidea based on the published articles from India.
The highest species richness (S = 18) was found in the Andaman and Nicobar Islands, followed by the East Coast (S = 13), West Coast (S = 12), and Lakshadweep (S = 8). The Andaman and Nicobar Islands also have the greatest Margalef diversity (d) at 3.638, followed by the West Coast at 3.07, the East Coast at 2.6, and Lakshadweep at 1.91. Pielou’s evenness (J’) evaluates both species richness and diversity. Pielou’s evenness was highest on the West Coast (0.86) and lowest in Lakshadweep (0.67). These findings show that the Andaman and Nicobar Islands have both the highest species richness and Margalef diversity, indicating a greater variety of species in this region compared to the others studied. Additionally, the West Coast exhibits high Pielou’s evenness, reflecting a more balanced distribution of species compared to the other three locations. By using the Bray–Curtis similarity measure, the degree of resemblance between data from existing documents can be quantified. The similarity indices of the four main stations are given in Figure 6. The results showed that the Andaman and Nicobar Islands, Lakshadweep, and the East Coast had a 40–60% level of resemblance. In comparison, the West Coast had very low (20%) similarities with the remaining regions.
Figure 6. Non-Metric Multidimensional scaling (nMDS) plot based on Bray–Curtis similarity patterns using data from the published studies from the West Coast, the East Coast, the Andaman and Nicobar Islands, and the Lakshadweep.

4. Discussion

This review has compiled checklist information on 187 species of holothuroids from the Indian waters, encompassing 7 orders and 21 families. The distribution of species within the class Holothuroidea in Indian waters is uneven, with the order Holothuriida (27%) having the highest percentage of species. This implies that specific ecological factors or environmental conditions may favor the abundance and diversity of holothuroids in this order.
The Andaman and Nicobar Islands showed a significant dominance of certain species and a higher diversity of holothuroids compared to other coastal regions included in this study. Asha et al. [42] made a similar observation in their study. Holothuroid habitats display a wide range of diversity, covering environments such as coral reefs, seagrass meadows, rocky shores, sandy beaches, muddy shores, and mangrove beds. The Andaman Islands are especially notable for hosting all these diverse habitats [82]. This variety of habitats may explain the greater abundance of holothuroids in this region compared to other parts of India. The East Coast and the Andaman Islands share a high similarity (upto 60%) in species distribution.
Notably, 101 species are reported on the East Coast, while 75 species are in the Gulf of Mannar. The Gulf of Mannar has an extensive range of echinoderms, due to its abundance of favorable habitats such as sandy bottoms, seagrass beds, and reefs [70]. The coral reefs and other macrofauna, particularly those found in the Gulf of Mannar and the islands of Andaman and Nicobar, are key reasons for the striking resemblance between these two locations [50]. The Andaman and Nicobar Islands and the West Coast, with their rich biodiversity and favorable environmental conditions, provide suitable habitats for many species, leading to higher diversity indices. Furthermore, [83] found that the Andaman and Nicobar Islands have high diversity, followed by the East Coast, West Coast, and Lakshadweep Islands. However, in this study, the West Coast showed a higher level of diversity compared to its eastern counterpart. This variation in diversity can be linked to the unique geographical features and ecosystems present in each area. In 2017, a checklist of echinoderms in India revealed a total of 179 holothuroids [81]. However, subsequent reclassifications have emerged, including the grouping of the families Synallactidae and Stichopodidae into the distinct order Synallactida. The order Aspidochorida is split into three orders, namely Persiculida, Synallactida [43], and Holothuriida. These reclassifications have resulted in changes in the classification of specific species. For instance, Stolus rapax has been reclassified as Pseudostolus rapax, while other species, such as Orphnurgus glaber and Oneirophanta conservata, have been reassigned to the order Synallactida [43].
In addition to the recent checklist, it is worth mentioning that the first sighting of Bohadschia atra in Indian waters was reported by Nithyanandan and Marimuthu [75], specimens collected from Kadmat and Agatti Islands in Lakshadweep. More recently, the same species has also been documented in the Andaman and Nicobar Islands [55]. Bohadschia atra is characterized by its brownish-black coloration, accentuated with red spots and numerous brown podia on its ventral side. Bhatt et al. [80] provided the first record of Holothuria (Microthele) fuscogilva occurrence along the Gujarat Coast. This species displays a beige or white color with yellowish undertones on its ventral side, while its dorsal surface is marked by black or dark brown mottling. These findings highlight the expanding distribution ranges of these species. Some species were misidentified and erroneously reported in previous publications [9,11,44,56]; for example, Apostichopus japonicus is restricted to the Northwest Pacific regions, and Parastichopus tremulus occurs only in the Northeast Atlantic and the deep Southwest Mediterranean Sea [12].
Numerous studies conducted in India have documented a gradual decline in both the diversity and density of sea cucumbers, largely driven by prolonged and unregulated exploitation. High-value species such as Holothuria scabra have experienced substantial reductions in population density, a trend attributed to continuous fishing pressure and limited capacity of depleted populations to recover [51,84]. Although all holothuroids are classified as protected, illegal harvesting persists in the Gulf of Mannar region, resulting in marked declines in key species such as Holothuria spinifera and H. scabra. Despite being aware of the illegality and the penalties associated with harvesting, local fishermen continue to engage in poaching [65]. The Andaman and Nicobar Islands, known for their rich marine biodiversity, have also experienced ongoing sea cucumber poaching [85]. Many poachers originate from neighboring countries, including Sri Lanka, Bangladesh, Myanmar, and Thailand, contributing directly or indirectly to the decline of sea cucumber populations in these regions [86]. A comprehensive study [87] revealed the illegal poaching of sea cucumber in the coast of Tamil Nadu and Andhra Pradesh. In our current study, the period from 1887 to 2024 indicates a consistent downward trend in sea cucumber abundance across Indian waters, primarily driven by over-exploitation and illegal harvesting. A similar observation was made by Choo [21] that overfishing is the primary cause of the decline in holothuroid resources across many Asian countries. Furthermore, the successful fertilization of subtropical and tropical sea cucumbers is strongly dependent on adequate population density; however, overfishing results in ineffective reproduction and severely limits population recovery [88].

5. Conclusions

The growing demand for holothuroids in the food industry has become a major concern, contributing significantly to the decline of their natural populations. Despite being listed as scheduled and protected species, sea cucumbers continue to be subject to intense exploitation driven by high commercial value, particularly in neighboring countries. Illegal poaching remains a widespread and persistent issue, posing a serious threat to the sustainability of holothuroid populations and the fragile marine ecosystems they support. The absence of an up-to-date species checklist further complicates efforts to accurately assess biodiversity within the Indian marine ecosystems. This lack of comprehensive information limits the development of effective conservation strategies and undermines responsible resource management. Moreover, it restricts the ability to identify species at risk of extinction that require immediate conservation action. Given these challenges, it is crucial to allocate resources and scientific expertise to regularly update species checklists to support evidence-based conservation in India. In this context, the present study provides the most recent and comprehensive checklist contains 187 species of the class Holothuroidea from Indian waters.

Author Contributions

Conceptualization, B.A.V.M.; Methodology, K.P.; Software, K.P.; Validation, B.A.V.M.; Formal analysis, K.P., A.U. and A.S.; Investigation, K.P., A.U. and A.S.; Data curation, K.P.; Writing—original draft, K.P.; Writing—review and editing, A.U., A.S. and B.A.V.M.; Visualization, B.A.V.M.; Supervision, B.A.V.M.; Funding acquisition, B.A.V.M. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The data presented in this study are available on request from the corresponding author (the data are not publicly available due to privacy restrictions).

Acknowledgments

The authors thank all reviewers for their constructive comments, which helped to improve the manuscript. K.P. thanks MEXT, Japan, for supporting with MEXT scholarship. B.A.V.M. thanks Organization for Marine Science and Technology, Nagasaki University, for the financial support.

Conflicts of Interest

The authors declare no conflicts of interest.

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