Hostname: page-component-74656f9884-npj9w Total loading time: 0 Render date: 2026-11-07T12:01:40.573Z Has data issue: false hasContentIssue false
  • English
  • Français

First report of the crinoid-associated scale worm Paradyte crinoidicola from the mainland Indian coastline

Published online by Cambridge University Press:  21 February 2025

Hitisha Baroliya Open the ORCID record for Hitisha Baroliya [Opens in a new window] ,
Jenish Kavathiya Open the ORCID record for Jenish Kavathiya [Opens in a new window]  and
Rahul Kundu Open the ORCID record for Rahul Kundu [Opens in a new window]
Hitisha Baroliya
Affiliation:
Department of Biosciences, Saurashtra University, Rajkot 360005, Gujarat, India
Jenish Kavathiya
Affiliation:
Department of Biosciences, Saurashtra University, Rajkot 360005, Gujarat, India
Rahul Kundu*
Affiliation:
Department of Biosciences, Saurashtra University, Rajkot 360005, Gujarat, India
*
Corresponding author: Rahul Kundu; Email: rskundu@sauuni.ac.in
Rights & Permissions [Opens in a new window]

Abstract

Feather stars (Echinodermata: Crinoidea) are known to host a diversified associated fauna. Surprisingly, very few accounts on the crinoid epiphytic communities from the coasts of the India are available. The present contribution reports for the first time the crinoid-associated scale worm Paradyte crinoidicola (Potts, 1910) from Gujarat coastline, India. The species has been so far recorded from the southern part of Indian subcontinent, the Lakshadweep Islands as its first record from Indian coastline. Further research and documentation of these species will contribute to enhance our understanding of the diverse associations and complex interactions within Indian marine ecosystems. This highlights the necessity for more thorough field surveys and ongoing monitoring, as the region remains surprisingly underexplored.

Keywords

feather starintertidalmainland Indian coastlinescale wormsymbiosis

Information

Type
Marine Record
Information
Journal of the Marine Biological Association of the United Kingdom , Volume 105 , 2025 , e25
Copyright
Copyright © The Author(s), 2025. Published by Cambridge University Press on behalf of Marine Biological Association of the United Kingdom

Introduction

Comatulid crinoids, or feather stars (Echinodermata: Crinoidea: Comatulida) are the most diversified order of living crinoids. Despite being distributed worldwide, they are key actors of coral reef benthic communities, where most of the extant species can be found (Messing and Tay, Reference Messing and Tay2016; Summers et al., Reference Summers, Messing and Rouse2017; Virgili et al., Reference Virgili, Poliseno, Fujita, Pratama, Fernández-Silva and Reimer2023). Feather stars are well-known hosts for a diverse associated community where various invertebrate and vertebrate organisms, such as annelids, crustaceans, gastropods and clingfish find food and shelter (e.g. Clark, Reference Clark1921; Zmarzly, Reference Zmarzly1984; Deheyn et al., Reference Deheyn, Lyskin and Eeckhaut2006; Britayev and Mekhova, Reference Britayev and Mekhova2011; Mekhova and Britayev, Reference Mekhova and Britayev2015; Britayev et al., Reference Britayev, Beksheneva, Deart and Mekhova2016; Virgili et al., Reference Virgili, Cerrano, Ponti, Lasut and Reimer2020). Among them, the phylum Annelida has one of the greatest diversities of associates with feather star (Britayev and Mekhova, Reference Britayev and Mekhova2011) with 180 species belonging to order Myzostomida and 7 species of order Phyllodocida (Martin and Britayev, Reference Martin and Britayev1998; Britayev et al., Reference Britayev, Doignon and Eeckahaut1999; Eeckhaut and Lanterbecq, Reference Eeckhaut, Lanterbecq, Bartholomaeus and Purschke2005; Britayev and Antokhina, Reference Britayev, Antokhina, Britayev and Pavlov2012; Summers et al., Reference Summers, Al-Hakim and Rouse2014). Investigations on the feather star-associated fauna in India has not attained much attention, and so far, brittle stars, crustaceans and myzostomids occurrence was recorded (Subramaniam, Reference Subramaniam1938; George, Reference George1943, Reference George1950; Rao and Sowbhagyavathi, Reference Rao and Sowbhagyavathi1974; Shyamasundari, Reference Shyamasundari1976; James, Reference James1995; Nigam and Raghunathan, Reference Nigam and Raghunathan2016; Mariyambi et al., Reference Mariyambi, Mohammednoushad, Idreesbabu and Sureshkumar2020; Prakash and Marimuthu, Reference Prakash and Marimuthu2020; Baroliya et al., Reference Baroliya, Solanki and Kundu2022). The associated polychaetae species, Paradyte crinoidicola (Potts, Reference Potts1910) is a generalist ectosymbiont with a wide distribution in the Indo-Pacific, adapted to live on a variety of crinoids hosts (Zmarzly, Reference Zmarzly1984; Deheyn et al., Reference Deheyn, Lyskin and Eeckhaut2006; Britayev et al., Reference Britayev, Beksheneva, Deart and Mekhova2016; Virgili et al., Reference Virgili, Cerrano, Ponti, Lasut and Reimer2020). The type locality of P. crinoidicola (Potts, Reference Potts1910) is Maldives Islands (Potts, Reference Potts1910). The species is widely distributed in the tropical Indo-West Pacific from Intertidal to 52 m depth (Britayev et al., Reference Britayev, Doignon and Eeckahaut1999). Recently, this species was reported for the first time from India, specifically from the subtidal areas of Lakshadweep Islands, in the southern Indian subcontinent (Mariyambi et al., Reference Mariyambi, Idreesbabu and Britayev2022). However, the true distribution of this species has never been evaluated from mainland India where records are missing. In the present study, we observed the scale worm, P. crinoidicola, in association with various comatulid crinoids from the intertidal areas of the Gujarat coast. It is also the first record of this species from the mainland Indian coastline.

Materials and methods

Gujarat is located at the western-most tip of the Indian subcontinent, has a 1600 km long stretch of coastline that includes three different areas, viz. Gulf of Kutch, Saurashtra coastline and Gulf of Khambhat. During the diversity survey targeting echinoderms, several scale worms were found associated with various crinoid species at Fort St. Anthony of Simbor (20.76305°N, 71.15634°E) and Nagao beach, Diu (20.16936°N, 71.82714°E), Gujarat, India (Figure 1). Fort St. Anthony of Simbor has a rocky outcrop with dense tide pools of feather stars (Baroliya and Kundu, Reference Baroliya and Kundu2021). There are also flat rocky structures with sandy-muddy substratum and pebbles which together provide a structural habitat for Crinoidea. We observed scale worms as symbiont of four crinoid species across both of the study sites. In the present study, the voucher specimens of crinoids with associated worms were collected by direct handpicking from the edges of the rock pool using soft paint brush. Collected crinoids with symbionts were placed to live in a 100 ml container filled with seawater and transferred to the laboratory. The live specimens were then photographed for their external morphology and colour pattern under the Lawrence & Mayo Image Stereo microscope (ISH300, BRAND). Identification of the scale worm and crinoids were based on the external morphological and taxonomical characters of the specimen using standard references (Potts, Reference Potts1910; Mariyambi et al., Reference Mariyambi, Idreesbabu and Britayev2022; crinoid: Clark, Reference Clark1921, Reference Clark1947; Pettibone, Reference Pettibone1969; Clark and Rowe, Reference Clark and Rowe1971; Messing, Reference Messing1997; Britayev et al., Reference Britayev, Doignon and Eeckahaut1999; Sastry, Reference Sastry2004; Britayev and Antokhina, Reference Britayev, Antokhina, Britayev and Pavlov2012; Mohammednowshad et al., Reference Mohammednowshad, Idreesbabu, Parameswaran, Messing and Sureshkumar2020; Baroliya et al., Reference Baroliya, Kavathiya and Kundu2024). The specimens were then anaesthetized using 7% MgCl2, preserved in 4% formalin and deposited in the museum of the Department of Biosciences, Saurashtra University, Rajkot, Gujarat, India. Museum ID: ZAP21(8)2H2, ZAP22(8)1H10, ZAP22(8)1H11, ZAP22(8)1H12.

Figure 1.

Map showing sampling localities of Paradyte crinoidicola (Potts, Reference Potts1910).

Results

Taxonomic account

Class Polychaeta Grube, 1850
Order Phyllodocida Dales, 1962
Family Polynoidae Kinberg, 1856
Genus Paradyte Pettibone, Reference Pettibone1969
Paradyte crinoidicola (Potts, Reference Potts1910)
(Fig. 3)
Polynoe crinoidicola Potts, Reference Potts1910: 337, pl. 18, figures 10, pl. 21, figures 39–41.
Scalisetosus (Polynoe) crinoidicola – Horst, Reference Horst1917: 98, pl. 16, figures 6–8, pl. 21, figure 1.
Adyte crinoidicola – Hanley, Reference Hanley1984: 88–90, figure 1.
Scalisetosus longicirrus Fauvel, Reference Fauvel and Sewell1953: 50, figures 22A–C.
Paradyte crinoidicola – Pettibone, Reference Pettibone1969: 13–16, figure 7; Tchesunov et al., Reference Tchesunov, Britayev, Larionov, Khodkina and Tzetlin1989: 169–172; Hanley and Burke, Reference Hanley, Burke and Crosnier1991: 13–15, figure 1; Britayev et al., Reference Britayev, Doignon and Eeckahaut1999: 369–371, figures 24–29; Barnich et al., Reference Barnich, Fiege and Sun2004, 312; Wehe, Reference Wehe2006: 154–156, figure 41; Britayev and Antokhina, Reference Britayev, Antokhina, Britayev and Pavlov2012: 22–23, pl. 4C–D; Mariyambi et al., Reference Mariyambi, Idreesbabu and Britayev2022: figures 2A–D.

Material examined

Fort St. Anthony Simbor: 1 specimen on Dichrometra palmata (Müller, 1841), 1 specimen on Tropiometra carinata (Lamarck, 1816), 2 specimens on Clarkcomanthus sp. (1 February 2021), 1 specimen on D. palmata (Müller, 1841), 1 specimen on T. carinata (Lamarck, 1816), 2 specimens on Comanthus wahlbergii (Müller, 1843), 3 specimens on Clarkcomanthus sp. (8 February 2022); Nagao Beach, Diu: 2 specimens on C. wahlbergii (Müller, 1843), 1 specimen on D. palmata (Müller, 1841) (9 February 2022), coll. H. Baroliya.

Description

Body elongated, fattened, tapered posteriorly, 12 mm long and 3.20 mm wide, 38 chaetigers and 15 pairs of elytra, posterior 5–6 chaetigers without elytra (Figure 2A). The prostomium is bilobed broader than long and with two pairs of eyes, minute frontal horns on the prostomial lobes, two palps and three antennae (Figure 2B). The elytra are smooth delicate, slightly rounded, lacking tubercles and sensory papillae (Figure 2C). Two transverse ciliary bands are present per segment. Notochaetae robust sabre-like, with few spines. Two forms of neurochaetae with semilunar pockets, supra-acicular cluster slender, slightly curved, with long spinous and bidentate tips, and sub-acicular are stouter, with absolute semicircular tips (Figure 2D).

Figure 2.

Morphological characteristics of Paradyte crinoidicola. (A) Whole specimen of P. crinoidicola. (B) Bilobed prostomium of P. crinoidicola. (C) Elytra of P. crinoidicola. (D) Parapodia (scale = A, B – 1 mm; C, D – 0.1 mm).

Notes on external coloration

The coloration is variable depending on the host, which can range from light pink to dark reddish brown. In the present study, individuals of P. crinoidicola collected from D. palmata have a dark stripe running longitudinally across the prostomium, and middle of the dorsal surface up to bases of parapodia while rest of the other individuals of P. crinoidicola with T. carinata, C. wahlbergii and Clarkcomanthus sp. were dark brown in colour (Figure 3).

Figure 3.

Plate shows host crinoids species with the presence of Paradyte crinoidicola indicated with circle. (A) Dichrometra palmata (Müller, 1841), (B, C) Clarkcomanthus sp., (D) Comanthus wahlbergii (Müller, 1843).

Ecological notes

In the present study crinoids were found attached with flat rocky substratum and shallow water tidepools, 0.2–0.5 m deep, rocky intertidal, Gujarat, India (Baroliya and Kundu, Reference Baroliya and Kundu2021). Paradyte crinoidicola was mostly found as an ectocommensal with the T. carinata, D. palmata, C. wahlbergii and Clarkcomanthus sp. In the present study, P. crinoidicola observed at the base of arms, hiding in arm's pinnules, on cirri and sometimes on the oral surface, typically resembles pinnules of the host species (Figure 3). Slight variation was observed in the colour, body comparison and elytra size. All the samples collected are complete and adult. No juvenile or damaged individuals were observed during the present study.

Discussion

The present study describes the crinoid-associated P. crinoidicola from the mainland of India. In the present study, P. crinoidicola was observed in association with four crinoid species, T. carinata, D. palmata, Clarkcomanthus sp. and C. wahlbergii, all from the intertidal zone of Gujarat, India. Each host harboured from one to three individuals. The aggregation of this worm was higher in Clarkcomanthus sp., probably related to the larger dichotomized arms of this species, providing more space and reducing intraspecific competition between symbionts (Britayev et al., Reference Britayev, Doignon and Eeckahaut1999). The present study is the first record about the association of polychaetas with echinoderms from the Gujarat coast and the whole mainland Indian coast. Further research and documentation of these species will contribute to enhance our understanding of the diverse associations and complex interactions within Indian marine ecosystems. Comparable studies on the crinoid-associated fauna suggest that the biodiversity of the Gujarat coastline could be higher than the current knowledge. This highlights the necessity for more thorough field surveys and ongoing monitoring, as the region remains surprisingly underexplored.

Acknowledgments

The authors are thankful for the laboratory facilities provided by the Department of Biosciences, Saurashtra University, Gujarat, India.

Author contributions

H. B. and R. K. conceived the study; H. B. and J. K. did the field work, collected data and identification; J. K. surveyed the literature; H. B. preceded the manuscripts writing, R. K. reviewed the manuscript. All authors contributed significantly to drafting the manuscript.

Financial support

The authors are thankful to the Department of Bioscience, Rajkot for supporting this study through its fieldwork fund provided by Saurashtra University.

Competing interest

The authors declare that they have no known competing interests that influence the work reported in this paper.

Ethical standards

All applicable international, national, and/or institutional guidelines for the care and use of animals have been followed by the authors.

Data availability statement

The authors confirm that the data supporting the findings of this study are available within the article itself.

References

Barnich, R, Fiege, D and Sun, R (2004) Polychaeta (Annelida) of Hainan Island, South China Sea Part III. Aphroditoidea. Species Diversity 9, 285–329.CrossRefGoogle Scholar
Baroliya, H, Kavathiya, J and Kundu, R (2024) First report of Tropiometra carinata and its colour morphs from the Gujarat coast, India. Spixiana 46, 165–171.Google Scholar
Baroliya, H and Kundu, R (2021) Intertidal aggregation of feather stars (Echinodermata: Crinoidea) on the south Saurashtra coastline of Gujarat state, India. International Journal of Ecology and Environmental Sciences 47, 339–344.Google Scholar
Baroliya, H, Solanki, B and Kundu, R (2022) Intertidal Ophiuroidea from the Saurashtra coastline, Gujarat, India. Journal of Threatened Taxa 14, 21968–21975.CrossRefGoogle Scholar
Britayev, TA and Antokhina, TI (2012) Symbiotic polychaetes from Nhatrang Bay, Vietnam. In Britayev, TA and Pavlov, DS (eds), Benthic Fauna of the Bay of Nhatrang, Southern Vietnam. Moscow: KMK , pp. 11–54.Google Scholar
Britayev, TA, Beksheneva, LF, Deart, YV and Mekhova, ES (2016) Structure and variability of symbiotic assemblages associated with feather stars (Crinoidea: Comatulida) Himerometra robustipinna. Oceanology 56, 666–674.CrossRefGoogle Scholar
Britayev, TA, Doignon, G and Eeckahaut, I (1999) Symbiotic polychaetes from Papua New Guinea associated with echinoderms, with descriptions of three new species. Cahiers de Biologie Marine 40, 359–374.Google Scholar
Britayev, TA and Mekhova, ES (2011) Assessment of hidden diversity of crinoids and their symbionts in the Bay of Nhatrang, Vietnam. Organisms Diversity & Evolution 11, 275.CrossRefGoogle Scholar
Clark, AH (1921) Monograph of the existing crinoids. the comatulids, part 2. Bulletin of the US National Museum 82, 1–795.Google Scholar
Clark, AH (1947) A monograph of the existing crinoids. 1(4b). Bulletin of the United States National Museum 82, 1–473, pl. 43.Google Scholar
Clark, AM and Rowe, FWE (1971) Monograph of shallow-water Indo-west Pacific Echinoderms. London: Trustees of the British Museum (Natural History), pp. 1–238.Google Scholar
Deheyn, DD, Lyskin, S and Eeckhaut, I (2006) Assemblages of symbionts in tropical shallow-water crinoids and assessment of symbionts’ host-specificity. Symbiosis 42, 161–168.Google Scholar
Eeckhaut, I and Lanterbecq, D (2005) Myzostomida: a review of their ultrastructure and phylogeny. In Bartholomaeus, T and Purschke, G (eds), Morphology, Molecules and Evolution of the Polychaeta and Related Taxa. The Netherlands: Springer, pp. 253–275.CrossRefGoogle Scholar
Fauvel, P (1953) Annelida Polychaeta. In Sewell, RS (ed.), The Fauna of India Including Pakistan, Ceylon, Burma and Malaya. Allahabad: The Indian Press, pp. 1–507.Google Scholar
George, AI (1943) On an undescribed species of Myzostoma from the Madras harbour with a note on the occurrence of Myzostoma adhaerens Remscheid. Proceeding of the Indian Science Congress 29, 150.Google Scholar
George, AI (1950) On Myzostoma striata, n. sp. from the Madras harbour. In Proceedings/Indian Academy of Sciences, New Delhi: Springer India 32(5), 215–221.Google Scholar
Hanley, JR (1984) A new host and locality records of the commensal Adyte crinoidicola (Polychaeta, Polynoidae). The Beagle, Records of the Northern Territory Museum of Arts and Sciences 1, 87–92.Google Scholar
Hanley, JR and Burke, M (1991) Polychaeta Polynoidae: scaleworms of the Chesterfeld Islands and Fairway Reefs, Coral Sea. In Crosnier, A (ed.), Résultats des Campaignes Musorstom, Vol. 8. Memory Museum of National History Nature (A) 151, 9–82. Paris.Google Scholar
Horst, R (1917) Polychaeta Errantia of the Siboga Expedition. Part 2. Aphroditidae and Chrysopetalidae. Siboga-Expeditie Uitkomsten op Zoologisch, Bonatisch. Oceanographisch en Geologisch gebied verzameld in Nederlandsch Oost-Indië 1899–1900 24b, 1–140.Google Scholar
James, DB (1995) Animal associations in echinoderms. Journal of the Marine Biological Association of India 37, 272–276.Google Scholar
Mariyambi, PC, Idreesbabu, KK and Britayev, TA (2022) First report of symbiotic polynoid scaleworm (Paradyte crinoidicola Potts, 1910) from the Indian subcontinent. Thalassas: An International Journal of Marine Sciences 38, 35–40.Google Scholar
Mariyambi, PC, Mohammednoushad, B, Idreesbabu, KK and Sureshkumar, S (2020) First record of two crinoid-associated Brachyuran crabs (Crustacea: Decapoda: Pilumnidae) from the Arabian Sea, Western Indian Ocean. Thalassas: An International Journal of Marine Sciences 36, 261–266.CrossRefGoogle Scholar
Martin, D and Britayev, TA (1998) Symbiotic polychaetes: review of known species. Oceanography and Marine Biology: An Annual Review 36, 217–340.Google Scholar
Mekhova, ES and Britayev, TA (2015) Soft substrate crinoids (Crinoidea: Comatulida) and their macrosymbionts in Halong Bay (North Vietnam). Raffles Bulletin of Zoology 63, 438–445.Google Scholar
Messing, CG (1997) Living comatulids. Paleontological Society Papers 3, 3–30.CrossRefGoogle Scholar
Messing, CG and Tay, TS (2016) Extant Crinoidea (Echinodermata) of Singapore. Raffles Bulletin of Zoology 34, 627–658.Google Scholar
Mohammednowshad, B, Idreesbabu, KK, Parameswaran, UV, Messing, CG and Sureshkumar, S (2020) The shallow-water crinoid fauna of Lakshadweep Atolls, north-western Indian Ocean. Zootaxa 4789, 247–265.CrossRefGoogle ScholarPubMed
Nigam, NK and Raghunathan, C (2016) New record of crinoid associated brittle star Gymnolophus obscura (Ljungman, 1967) from Andaman and Nicobar Islands, India. World Journal of Fish and Marine Sciences 8, 06–09.Google Scholar
Pettibone, MH (1969) Review of some species referred to Scalisetosus McIntosh (Polychaeta, Polynoidae). Proceedings of the Biological Society of Washington 82, 1–30.Google Scholar
Potts, FA (1910) Polychaeta of the Indian Ocean. Pt. 2. The Palmyridae, Aphroditidae, Polynoidae, Acoetidae and Sigalionidae. Transactions of the Linnean Society of London 16, 325–353.CrossRefGoogle Scholar
Prakash, S and Marimuthu, N (2020) Notes on some crinoid associated decapod crustaceans (Crustacea: Decapoda) of Lakshadweep Archipelago, Central Indian Ocean. Zootaxa 4766, 86–100.CrossRefGoogle ScholarPubMed
Rao, KH and Sowbhagyavathi, R (1974) Observations on the associates of crinoids at Waltair Coast with special reference to myzostomes. Proceedings of the National Academy of Sciences, India, Section B: Biological Sciences 38, 360–366.Google Scholar
Sastry, DRK (2004) Echinodermata: Crinoidea, Asteroidea, Ophiuroidea and Echinoidea. Fauna of Gujarat Part-2 State fauna series 8, 1–427.Google Scholar
Shyamasundari, K (1976) Comparative cytochemistry of female gametes of invertebrates with special reference to Myzostoma gopalai Subramanyam, 1938 and Myzostomum striatum George, 1950 (Polychaeta, Myzostomida). Acta Parasitologica Polonica 24, 199–202.Google Scholar
Subramaniam, MK (1938) On Myzostoma gopalai species nova from the Madras harbour. In Proceedings/Indian Academy of Sciences, New Delhi: Springer India 7(5), 270–276.Google Scholar
Summers, MM, Al-Hakim, II and Rouse, GW (2014) Turbo-taxonomy: 21 new species of Myzostomida (Annelida). Zootaxa 3873, 301–344.CrossRefGoogle ScholarPubMed
Summers, MM, Messing, CG and Rouse, GW (2017) The genera and species of Comatulidae (Comatulida: Crinoidea): taxonomic revisions and a molecular and morphological guide. Zootaxa 4268, 151–190.CrossRefGoogle Scholar
Tchesunov, AV, Britayev, TA, Larionov, VV, Khodkina, IV and Tzetlin, AB (1989) Notes on commensals of some crinoids from Maldives coral reef. In: Symbiosis in Marine Animals. Sveshnikov, V.A., ed. Vaskhnil, Moscow pp 166192.Google Scholar
Virgili, R, Cerrano, C, Ponti, M, Lasut, MT and Reimer, JD (2020) Crinoid diversity and their symbiotic communities at Bangka Island (North Sulawesi, Indonesia). Marine Biodiversity 50, 1–24.CrossRefGoogle Scholar
Virgili, R, Poliseno, A, Fujita, T, Pratama, GA, Fernández-Silva, I and Reimer, JD (2023) Molecular phylogeny of the genus Dorometra Clark, 1917 (Crinoidea: Comatulida: Antedonidae): A new genus and new insights for future taxonomic revisions of Antedonidae. Systematic & Biodiversity 21, 2192209.CrossRefGoogle Scholar
Wehe, T (2006) Revision of the scaleworms (Polychaeta: Aphroditoidea) occurring in the seas surrounding the Arabian Peninsula. Part I: Polynoidae Fauna Arab 22, 23–197.Google Scholar
Zmarzly, DL (1984) Distribution and ecology of shallow-water crinoids at Enewetak Atoll, Marshall Islands, with an annotated checklist of their symbionts. Pacific Science 38, 105–122.Google Scholar
Figure 0

Figure 1. Map showing sampling localities of Paradyte crinoidicola (Potts, 1910).

Figure 1

Figure 2. Morphological characteristics of Paradyte crinoidicola. (A) Whole specimen of P. crinoidicola. (B) Bilobed prostomium of P. crinoidicola. (C) Elytra of P. crinoidicola. (D) Parapodia (scale = A, B – 1 mm; C, D – 0.1 mm).

Figure 2

Figure 3. Plate shows host crinoids species with the presence of Paradyte crinoidicola indicated with circle. (A) Dichrometra palmata (Müller, 1841), (B, C) Clarkcomanthus sp., (D) Comanthus wahlbergii (Müller, 1843).