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Amphipholis squamata (Delle Chiaje 1828) Asteria squamata Delle Chiaje, 1828: 74. Amphipholis squamata.— Mortensen 1933a: 364–365.—Clark, A.M. 1976: 258.— Olbers et al. 2019: 217–218, fig. 214–215. Ophiactis minor Döderlein, 1910: 253, pl. 5(3–3a) [see Mortensen 1933b]. Amphipholis minor.— Hertz 1927a: 35. Material examined. MD 50 DC2, MNHN IE.2009.1522 (2). MD 50 WP4, MNHN IE.2009.1523 (1). MD 50 DC8, MNHN IE.2009.1524 (5). MD 50 DC44, MNHN IE.2009.1525 (16). MD 50 DC59, MNHN IE.2009.1526 (4). MD 50 DC71, MNHN IE.2009.1527 (2). MD 50 DC79, MNHN IE.2009.1528 (2). MD 50 DC105, MNHN IE.2009.1529 (1). MD 50 DC114, MNHN IE.2009.1530 (1). MD 50 CP149, MNHN IE.2009.1531 (9). MD 50 DC157, MNHN IE.2009.1532 (21). MD 50 DC167, MNHN IE.2009.1533 (1). Distribution. Arctic (29–2369 m), NW Atlantic (0–1962 m), NE Atlantic (0–1560 m), NW Pacific (0–413 m), NE Pacific (0–933 m), W Atlantic (0–353 m), E Atlantic (0–741 m), W Indian (0–750 m), E Indo-W Pacific (0–694 m), E Pacific (0–46 m), S America (0–134 m), S Africa (0–1600 m), S Australia (0–841 m), New Zealand (2–1059 m). SPA (3–1600 m). Remarks. Amphipholis squamata is a polyphyletic (O’Hara et al. 2017) assemblage of numerous genetic lineages (Boissin et al. 2008). Only synonyms relevant to the SPA are listed above. The wide bathymetric range of the MD50 samples suggests that several lineages may be present in the SPA. Hertz (1927a) named her specimens A. minor after a South African lineage, however, the diagnostic characters used by Döderlein (1910) to distinguish this species were inadequate (Mortensen 1933a). The MD50 material is small and damaged., Published as part of O'Hara, Timothy D. & Thuy, Ben, 2022, Biogeography and taxonomy of Ophiuroidea (Echinodermata) from the Îles Saint- Paul and Amsterdam in the southern Indian Ocean, pp. 1-49 in Zootaxa 5124 (1) on pages 38-39, DOI: 10.11646/zootaxa.5124.1.1, http://zenodo.org/record/6404674, {"references":["Delle Chiaje, S. (1828) Memoire sulla Storia e Notomia degli Animali senza vertebre del Regno di Napoli. Vol. 3. Societa Tipografica, Napoli, 232 pp.","Mortensen, T. (1933 a) Echinoderms of South Africa (Asteroidea: Ophiuroidea). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 93, 215 - 400, pls. 8 - 19.","Olbers, J. M., Griffiths, C. L., O'Hara, T. D. & Samyn, Y. (2019) Field guide to the brittle and basket stars (Echinodermata: Ophiuroidea) of South Africa. Brussels: Royal Belgium Institute of Natural Sciences.","Doderlein, L. (1910) Asteroidea, Ophiuroidea, Echinoidea. In: Schultze, L. (Ed.), Forschungsreise im westlichen Sudafrica 4 (1). Denkschriften der Medizinisch-Naturwissenschaftlichen Gesellschaft zu Jena, 16, pp. 245 - 258, pls. 4 - 5.","Mortensen, T. (1933 b) The echinoderms of St. Helena. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i KObenhavn, 93, 401 - 472.","Hertz, M. (1927 a) Die Ophiuroiden der Deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition, 19, 1 - 56, pls. 1 - 9.","O'Hara, T. D., Hugall, A. F., Thuy, B., Stohr, S. & Martynov, A. V. (2017) Restructuring higher taxonomy using broad-scale phylogenomics: the living Ophiuroidea. Molecular Phylogenetics and Evolution, 107, 415 - 430. https: // doi. org / 10.1016 / j. ympev. 2016.12.006","Boissin, E., Feral, J. P. & Chenuil, A. (2008) Defining reproductively isolated units in a cryptic and syntopic species complex using mitochondrial and nuclear markers: the brooding brittle star, Amphipholis squamata (Ophiuroidea). Molecular Ecology, 17, 1732 - 1744. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2007.03652. x"]}

Amphipholis squamata (Delle Chiaje, 1828) Asterias squamata Delle Chiaje, 1828: 74, pl. 34, fig. 1. Amphipholis squamata — Ljungman 1872: 633. Material examined: Gulf of Oman, Iran, Tis, intertidal, hand collected, January 2019, collector Yaser Fatemi, 1 spm [not kept]. Habitat: Intertidal shell and gravel bed, under a stone (present study), grass bed and subtidal sand, 10–34 m (Price 1981; present study) (Fig. 2E). Persian Gulf and Gulf of Oman: Tarut Bay (25), Jinna Island (21), Tis (16), Bahrain (ibid.). Indian Ocean: Southern Africa (Clark & Courtman-Stock 1976), West Indian Ocean (Clark & Rowe 1971), Gulf of Mannar (James 1985), Pakistan (Kazmi & Tahera 2003), Mascarene Isl. (Boissin et al. 2016), Scattered Isl. (Conand et al. 2016). Remarks: A record from Chabahar Bay by Attaran-Fariman & Beygmoradi (2016) is an error and omitted here. Their images show an animal with narrow, sunken in radial shields (Amphipholis has short, half-circle shaped ones), bell-shaped overlapping dorsal arm plates (separated in A. squamata), small narrow tentacle scales (larger oval in A. squamata), widely separated infradental papillae (close together in A. squamata), and the distal oral papilla is only slightly widened, not as wide as in A. squamata. In addition, according to the scale bar on the images, the animal had a disc diameter of 8 mm, which is larger than A. squamata (maximum 5 mm), and they reported the arms to be 6 cm long, whereas A. squamata has about 2 cm long arms. The characters fit with Amphiodia, but the images do not show the necessary details for species identification. Consequently, the material found by the present study is the first record of A. squamata from the Gulf of Oman. Amphipholis squamata has a circumtropical distribution, but it is most likely a species complex (Sponer & Roy 2002; Boissin et al. 2015)., Published as part of Fatemi, Yaser & Stöhr, Sabine, 2019, Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records, pp. 77-106 in Zootaxa 4711 (1) on page 99, DOI: 10.11646/zootaxa.4711.1.3, http://zenodo.org/record/3573466, {"references":["Delle Chiaje, S. (1828) Memorie sulla storia e anotomia degli animali senza vertebre del Regno di Napoli. Vol. 3. Stamperia della Societa Tipografica, Naples, 232 pp.","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskapsakademiens Forhandlingar 1871, 6, 615 - 658.","Price, A. R. G. (1981) Studies on the echinoderm fauna of the western Arabian Gulf. Journal of Natural History, 15, 1 - 15. https: // doi. org / 10.1080 / 00222938100770011","Clark, A. M. & Courtman-Stock, J. (1976) The echinoderms of Southern Africa. British Museum, London, 277 pp.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 234 pp.","James, D. B. (1985) Echinoderm fauna of the proposed National Marine Park in the Gulf of Mannar. Proceeding of Symposium Endangered Marine Animals and Marine Parks, 1, 403 - 406.","Kazmi, Q. & Tahera, Q. (2003) A new record of Amphipholis squamata (Delle Chiaje, 1828) (Echinodermata: Ophiuroidea) in Pakistan waters. Pakistan Journal of marine science, 12, 91 - 92.","Boissin, E., Hoareau, T. B., Paulay, G. & Bruggemann, J. H. (2016) Shallow-water reef ophiuroids (Echinodermata: Ophiuroidea) of Reunion (Mascarene Islands), with biogeographic considerations. Zootaxa, 4098 (2), 273 - 297. https: // doi. org / 10.11646 / zootaxa. 4098.2.4","Conand, C., Mulochau, T., Stohr, S., Eleaume, M. & Chabanet, P. (2016) Inventory of echinoderms in the Iles Eparses (Europa, Glorieuses, Juan de Nova), Mozambique Channel, France. Acta Oecologica, 72, 53 - 61. https: // doi. org / 10.1016 / j. actao. 2015.06.007","Attaran-Fariman, G. & Beygmoradi, A. (2016) Short communication: The first report of Amphipholis squamata (Delle Chiaje, 1829) (Echinodermata: Ophiuroidea) from Chabahar Bay - northern Oman Sea. Iranian Journal of Fisheries Sciences, 15, 1254 - 1261.","Sponer, R. & Roy, M. S. (2002) Phylogeographic analysis of the brooding brittle star Amphipholis squamata (Echinodermata) along the coast of New Zealand reveals high cryptic variation and cryptic dispersal potential. Evolution, 56, 1954 - 1967.","Boissin, E., Egea, E., Feral, J. P. & Chenuil, A. (2015) Contrasting population genetic structures in Amphipholis squamata, a complex of brooding, self-reproducing sister species sharing life history traits. Marine Ecology Progress Series, 539, 165 - 177. https: // doi. org / 10.3354 / meps 11480"]}

Artisanal small-scale gold mining (ASGM) using mercury (Hg) amalgamation commenced on Buru Island, Indonesia, in 2012, but was halted in 2015 due to concerns of widespread Hg contamination. Much of the Hg used in the mining process is lost to trommel waste which is disposed of in settlement ponds that drain into adjacent waterways and into Kayeli Bay. Several thousand unmanaged trommel sites and associated tailing ponds exist on Buru Island. This study shows that waste from the Marloso trommel at the Gogrea site contained 203 mg/kg total Hg (THg), with a negligible proportion present as bioavailable methyl Hg (MeHg) and a low total organic carbon content. There are currently very few tools available for ecotoxicological risk assessment of mine tailings for tropical marine ecosystems, and we support the development of Tailings Toxicity Tests (TTTs) and describe laboratory toxicity test methods using the cosmopolitan benthic echinoderm Amphipholis squamata. Undiluted trommel waste caused 100% mortality of A. squamata within 48 h, and a 96-h LC50 of 6.7% w/w trommel waste (4 mg/kg THg) was estimated. Sub-lethal effects on the water vascular system of the brittle star were assessed by quantification of the Ability to Right Itself (ARI), and a 48-h EC50 of 7.3% w/w trommel waste (14.4 mg/kg THg) was estimated. The results show that trommel waste produced on Buru Island is highly contaminated with THg and is acutely toxic, raising serious concern for receiving ecosystems where Hg methylation to more toxic and bioavailable forms is likely.

The fine structure of the male reproductive system of the hermaphroditic brittle-star, Amphipholis squamata, has been studied in specimens from both the Pacific coast (Washington) and the Atlantic coast (New Hampshire). Each testis is a small (100-μm) sphere and is attached to the internal wall of the bursa by peritoneal suspensor cells. Occasional flagellated cells are found on the external surface of the testis. The testicular wall of A. squamata is a multilayered structure, similar to that of other ophiuroids, but the hemal sinus is PAS negative in this species. Germinal cells are surrounded throughout their development by the filopodia of interstitial cells. Adjacent interstitial cells are interconnected, and thus form a structural network within the testis. Positionally and functionally, the interstitial cells resemble Sertoli cells; however, their origin, behavior and ultrastructure are different in many ways. Spermatogenesis includes a series of cyclical changes (aspermatogenic phase, proliferative phase, differentiative phase, and evacuative phase). Within a single testis, the resulting production of sperm is in short pulses, but if all 10 testes are taken together, sperm are produced continuously throughout the year. The events of spermiogenesis closely follow those that have been described in other echinoderms. However, we have provided new information on the release of excess cell membranes and the fusion process of mitochondria. The mature spermatozoa of A. squamata are flagellated and motile, and have “primitive” structural features, in spite of the fact that they fertilize the eggs inside the genital bursae. The spermatozoa do not, as was previously thought, enter the bursa by rupture of the adjacent walls. Instead, they are ejaculated through a gonoduct into the rapid incurrent flow of water entering the bursa. The locomotion of the spermatozoa is in eccentric spirals, due to the unusually large angle at which the flagellum is inserted into the base of the sperm.

Amphipholis squamata (Delle Chiaje, 1828) (Fig. 15) Type locality. Mediterranean Sea. Maximum size. dd up to 5.5 mm (present study). Material examined. 53 specimens (dd: 2.4–5.5 mm). Intertidal and subtidal: ZUEC OPH 2121, St. 26, 2 spms; ZUEC OPH 2122, St. III, 1 spm; ZUEC OPH 2124, St. 32, 1 spm; ZUEC OPH 2158, St. VI, 1 spm; ZUEC OPH 2179, St. XII, 1 spm; ZUEC OPH 2187, St. XI, 7 spms; ZUEC OPH 2201, St. 68, 1 spm; ZUEC OPH 2209, St. 71, 1 spm; ZUEC OPH 2250, St. XIX, 2 spms; ZUEC OPH 2293, St. 119, 1 spm; ZUEC OPH 2330, St. 9H, 1 spm; ZUEC OPH 2331, St. 10H, 1 spm; ZUEC OPH 2336, St. 11H, 1 spm; ZUEC OPH 2456, St. 16H, 1 spm; ZUEC OPH 2457, St. 22H, 1 spm. From rocky shore in sponge: ZUEC OPH 2150, St. 3C, 2 spm; ZUEC OPH 2151, St. 5C, 1 spm; ZUEC OPH 2152, St. 7C, 1 spm; ZUEC OPH 2154, St. 9C, 9 spms; ZUEC OPH 2429, St. 11C, 3 spms; ZUEC OPH 2431, St. 11C, 1 spm; ZUEC OPH 2436, St. 13C, 2 spm; ZUEC OPH 2454, St. 15C, 10 spms; ZUEC OPH 2458, St. 15C, 1 spm. Description. Disc: (dd: 3.5 mm) circular, covered by irregular and imbricated scales, approximately 13 between the centrodorsal and the edge of the disc. Central primary plates and primary radial plates evident, larger than the scales. Radial shields twice as long as wide, contiguous for almost their entire length, separated proximally by one triangular scale (Fig. 15A). Ventral interradius covered with scales smaller than the dorsal ones and strongly imbricated. Bursal slits long and wide (Fig. 15B). Oral shields diamond-shaped, as long as wide. Madreporite larger than other oral shields and with rounded edges. Adoral shields well developed, broadened distally and united proximally. Two lateral oral papillae, distal rectangular and wider than the proximal one. A pair of elongated infradental papillae (Fig. 15C). Arms: dorsal arm plates broadly oval and contiguous (Fig. 15D,F). Ventral arm plates pentagonal, twice as long as wide with lateral edges concave near the tentacle pores, proximal angle pointed and distal edge straight, contiguous (Fig. 15E,G). Two subequal tentacle scales, one inserted on the ventral arm plate and the other on the lateral arm plate (Fig. 15E). Three pointed arm spines (Fig. 15D). Lateral arm plates (Fig. 15H,I): general outline: ventral portion projecting ventro-proximalwards; ventro-distal not projecting ventralwards. Outer surface ornamentation: trabecular intersections protruding to form knobs approximately the same size than stereom pores. Outer proximal edge: surface lined by discernible band of different stereom structure, but only in central part; without spurs; central part not protruding; surface without horizontal striation. Spine articulations: on same level as remaining outer surface, sizes all similar; distance between spine articulation equidistant. Lobes simply separated, equal-sized; lobes parallel, straight, and lobes oriented nearly horizontal; stereom massive; sigmoidal fold absent. Inner side, ridges and knobs: dominated by two separate central knobs; without additional dorsal structure on inner side; single large perforation on inner side. Vertebrae: zygospondylous of universal type and non-keeled. Proximal side of vertebrae dorsally without large groove on the dorsal-distal muscular fossae (Fig. 15J). Zygocondyles dorsalwards converging and zygosphene fused with pair of zygocondyles (Fig. 15K). Dorso-distal muscular fossae transformed distalwards projecting but far from distal edge of zygocondyles and with a vertical oval hole through the ossicle, about half as long as the vertebra (Fig. 15L,M). Zygosphene projecting beyond ventral edge of zygocondyles with projecting part longer than zygocondyles (Fig. 15M). Taxonomic comments. One unusually large specimen with 5.5 mm dd was identified (mean range 2.5–3.5 mm dd; Gondim et al. 2013a; Paim et al. 2015). A. squamata has a broad geographic distribution and forms a cryptic species complex (Sponer & Roy 2002; Boisson et al. 2008). Studies have failed to find any diagnostic characters, but they are suspected to be in reproductive isolation (Sponer et al. 2001; Boissin et al. 2010). Although genetic markers appear to be appropriate tool to delineate species of the A. squamata complex, (Sponer & Roy 2002; Le Gac et al. 2004; Boissin et al. 2008; Boissin et al. 2010), there are no detailed morphological studies with a broad geographic scale published. Its taxonomy should continue to be investigated using additional tools, such as description of arm ossicles. Consequently, a revision of its geographic distribution will be necessary (Rodrigues et al. 2011). Remarks. It produces bioluminescence only in the arms which is attributed to specific photocytes under ganglial control (Deheyn et al. 2000). This species can switch from deposit feeding by collecting particles with its tube feet to suspension feeding via trapping detritus in mucus (Rodrigues et al. 2011). A. squamata dwells on a variety of substrates including sand and rocky bottom as well as biological substrates such as sponges, bryozoans, corals, molluscs, polychaetes, and seagrass (Borges & Amaral 2005; Gondim et al. 2013a; Paim et al. 2015). It was collected from the sponge Amphimedon viridis (44% of spms), sand (fine, medium and coarse) and rubble bottom with a dredge (36% of spms) and van Veen grab (20% of spms). Distribution. Wide distribution, absent only from the Polar Regions (Hendler et al. 1995). In Brazil, it has been recorded from the Tropical Atlantic (realm), Tropical Southwestern Atlantic (province): Northeastern Brazil (Lima-Verde 1969; Magalhães et al. 2005; Gondim et al. 2008; Manso et al. 2008; Lima et al. 2011; Gondim et al. 2013a; Gondim et al. 2013b; Paim et al. 2015), Trindade and Martin Vaz Islands (Tommasi & Aron 1987), to Temperate South America (realm), Warm Temperate Southwestern Atlantic (province): Southeastern Brazil (Clark 1915; Manso & Absalão 1988; Pires-Vanin et al. 1997; Hadel et al. 1999; Borges et al. 2002; Netto et al. 2005; Pires-Vanin et al. 2014). From intertidal to 1200 m depth (Alvarado & Solís-Marín 2013). The present study samples occurred at depths ranging from intertidal to 21.5 m. Selected references. Delle Chiaje (1828): p. 74, fig. 1–4 [as Asterias squamata]; Ljungman (1872): p. 647 [as Amphipholis appressa]; Clark (1909): p. 540, fig. lii 1–3 [as Amphipholis australiana]; (Farquhar, 1897) Ljungman (1867): p. 312; Ljungman (1872): p. 646 [as Amphipholis elegans]; Matsumoto (1915): p. 71; Matsumoto (1917): p. 186, fig. 49 [as Amphipholis japonica]; Ljungman (1872): p. 646 [as Amphipholis kinbergi]; Ljungman (1872): p. 645 [as Amphipholis lineata]; Hertz (1927): p. 35 [as Amphipholis minor]; Ljungman (1872): p. 646 [as Amphipholis patagonica]; Thomas (1962): p. 662, fig. 13; Madsen (1970): p. 202, fig. 30; Paterson (1985): p. 91, fig. 36; Hendler et al. (1995): p. 162, fig. 79; Borges et al. (2002): p. 44, fig. 24a–b; Borges & Amaral (2005): p. 261, fig. a–d; Manso et al. (2008): p. 191, fig. 19a–c; Gondim et al. (2013a): p. 61, fig. 5 g –l; Paim et al. (2015): p. 5, fig. 2d–f [as Amphipholis squamata]; Ljungman (1865): pl. XV [as Amphipholis squamata tenuispina]; Ljungman (1872): p. 634 [as Amphipholis tenera]; (Ljungman, 1865): Ljungman (1872): p. 633 [as Amphipholis tenuispina]; Forbes (1843): 150 [as Amphiura neglecta]; Hutton (1878): p. 305; Clark (1915): p. 230, pl. 5 fig. 10, 11 [as Amphiura parva]; Lyman (1865): p. 121 [as Amphiura squamata]; Lyman (1865): p. 123; Rathbun (1879): p. 154 [as Amphiura tenera]; Thomas (1966): p. 831; Tommasi (1970): p. 37, fig. 36 [as Axiognathus squamata]; Döderlein (1910): p. 253, Taf. V, fig. 3–3a [as Ophiactis minor]; Ayres (1852): p. 133 [as Ophiolepis tenuis]; Leach et al. (1815): p. 57 [as Ophiura elegans]., Published as part of Alitto, Renata A. S., Bueno, Maristela L., Guilherme, Pablo D. B., Domenico, Maikon Di, Christensen, Ana Beardsley & Borges, Michela, 2018, Shallow-water brittle stars (Echinodermata: Ophiuroidea) from Araçá Bay (Southeastern Brazil), with spatial distribution considerations, pp. 1-66 in Zootaxa 4405 (1) on pages 43-45, DOI: 10.11646/zootaxa.4405.1.1, http://zenodo.org/record/3097993, {"references":["Delle Chiaje, S. (1828) Memorie sulla storia e notomia degli animali senza vertebre del regno di Napoli. Napoli: Fratelli Fernandes, and Societa Tipografica, 1, 1 - 232. https: // doi. org / 10.5962 / bhl. title. 10021","Gondim, A. I., Alonso, C., Dias, T. L. P., Manso, C. L. C. & Christoffersen, M. L. (2013 a) A taxonomic guide to the brittle-stars (Echinodermata, Ophiuroidea) from the State of Paraiba continental shelf, Northeastern Brazil. ZooKeys, 307, 45 - 96. https: // doi. org / 10.3897 / zookeys. 307.4673","Paim, F. G., Guerrazzi, M. C. & Borges, M. (2015) Descriptive and illustrated diagnosis of Ophiuroidea fauna (Echinodermata) in the shallow waters of North-eastern Brazil. Marine Biodiversity Records, 8, 1 - 17. https: // doi. org / 10.1017 / S 1755267215000652","Sponer, R. & Roy, M. S. (2002) Phylogeographic analysis of the brooding brittle star Amphipholis squamata (Echinodermata) along the coast of New Zealand reveals high cryptic genetic variation and cryptic dispersal potential. Evolution, 56, 1954 - 1967. https: // doi. org / 10.1111 / j. 0014 - 3820.2002. tb 00121. x","Sponer, R., Deheyn, D. & Roy, M. S. (2001) Large genetic distances within a population of Amphipholis squamata (Echinodermata; Ophiuroidea) do not support colour varieties as sibling species. Marine Ecology Progress Series, 219, 169 - 175. https: // doi. org / 10.3354 / meps 219169","Boissin, E., Chenuil, A. & Feral, J. - P. (2010) Species of the complex Amphipholis squamata (Ophiuroidea) from Marseilles. In: Harris, L. G., Boetger, A. S., Walker, C. W. & Lesser, M. P. (Eds.), Echinoderms: Durham. Balkema, Rotterdam, Durham, New Hampshire, pp. 135 - 138. https: // doi. org / 10.1201 / 9780203869543 - c 20","Le Gac, M., Feral, J. - P., Poulin, E., Veyret, M. & Chenuil, A. (2004) Identification of allopatric clades in the cosmopolitan ophiuroid species complex Amphipholis squamata (Echinodermata). The end of a paradox? Marine Ecology Progress Series, 278, 171 - 178. https: // doi. org / 10.3354 / meps 278171","Boissin, E., Feral, J. - P. & Chenuil, A. (2008) Defining reproductively isolated units in a cryptic and syntopic species complex using mitochondrial and nuclear markers: the brooding brittle star, Amphipholis squamata (Ophiuroidea). Molecular Ecology, 17, 1732 - 1744. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2007.03652. x","Rodrigues, C. F., Paterson, G. L. J., Cabrinovic, A. & Cunha, M. R. (2011) Deep-sea ophiuroids (Echinodermata: Ophiuroidea: Ophiurida) from the Gulf of Cadiz (NE Atlantic). Zootaxa, 2754, 1 - 26.","Deheyn, D., Mallefet, J. & Jangoux, M. (2000) Cytological changes during bioluminescence production in dissociated photocytes from the ophiuroid Amphipholis squamata (Echinodermata). Cell and tissue research, 299, 115 - 128. https: // doi. org / 10.1007 / s 004419900144","Borges, M. & Amaral, A. C. Z. (2005) Classe Ophiuroidea. In: Amaral, A. C. Z., Rizzo, A. E. & Arruda, E. P. (Eds.), Manual de identificacao dos invertebrados marinhos da regiao sudeste-sul do Brasil. EdUSP, Sao Paulo, pp. 237 - 272.","Hendler, G., Miller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea stars, sea urchins, and allies: echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp. https: // doi. org / 10.1017 / s 0025315400031568","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do nordeste brasileiro. Arquivos de Ciencias do Mar, 9, 9 - 13.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9, 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Gondim, A. I., Lacouth, P., Alonso, C. & Manso, C. L. C. (2008) Echinodermata da Praia do Cabo Branco, Joao Pessoa, Paraiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8, 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Lima, M. L. F., Correia, M. D., Sovierzoski, H. H. & Manso, C. L. C. (2011) New records of Ophiuroidea (Echinodermata) from shallow waters off Maceio, State of Alagoas, Brazil. Marine Biodiversity Records, 4, 1 - 10. https: // doi. org / 10.1017 / S 175526721100090 X","Gondim, A. I., Dias, T. L. P. & Christoffersen, M. L. (2013 b) Annotated checklist of Echinoderms from Maranhao and Piaui States, Northeastern Brazil. Check List, 9, 510 - 518. https: // doi. org / 10.15560 / 9.3.510","Tommasi, L. R. & Aron, M. A. (1987) Equinodermes dos bancos submarinos da cadeia de montanhas Vitoria-Trindade. Relatorios Internos do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 9.","Clark, H. L. (1915) Catalogue of Recent Ophiurans: Based on the Collection of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology, 25, 165 - 376.","Manso, C. L. C. & Absalao, R. S. (1988) Ophiuroidea: Situacao Pre Operacional nos sacos de Piraquara, regiao sob influencia da descarga da Central Nuclear Almirante Alvaro Alberto (CNAAA). Revista Brasileira de Biologia, 48, 75 - 82.","Pires-Vanin, A. M. S., Corbisier, T. N., Arasaki, E. & Moellmann, A. M. (1997) Composicao e distribuicao espaco-temporal da fauna bentica no Canal de Sao Sebastiao. Relatorios Tecnicos do Instituto Oceanografico, 41, 29 - 46.","Hadel, V. F., Monteiro, A. M. G., Ditadi, A. S. F., Tiago, C. G. & Tommasi, L. R. (1999) Echinodermata. In: Migotto, A. E. & Tiago, C. G. (Eds.), Biodiversidade do Estado de Sao Paulo, Brasil: sintese do conhecimento ao final do seculo XX. Fundacao de Amparo a Pesquisa do Estado de Sao Paulo, Sao Paulo, pp. 260 - 271.","Borges, M., Monteiro, A. M. G. & Amaral, A. C. Z. (2002) Taxonomy of Ophiuroidea (Echinodermata) from the continental shelf and slope of the southern and southeastern Brazilian coast. Biota Neotropica, 2, 1 - 69. https: // doi. org / 10.1590 / S 1676 - 06032002000200010","Netto, L. F., Hadel, V. F. & Tiago, C. G. (2005) Echinodermata from Sao Sebastiao Channel (Sao Paulo, Brazil). Revista de Biologia Tropical, 53, 207 - 218.","Pires-Vanin, A. M. S., Muniz, P. & Bromberg, S. (2014) Inventory of the marine soft bottom macrofauna of Sao Sebastiao Channel, southeastern Brazilian continental shelf. Check List, 10, 795 - 807. https: // doi. org / 10.15560 / 10.4.795","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Springer, Berlin / Heidelberg, 665 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Ljungman, A. V. (1872) Forteckning ofver uti Vestindien af Dr A. Goes samt under korvetten Josefinas expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af Kungliga Vetenskaps-Akademiens Forhandlingar, 28, 615 - 658.","Clark, H. L. (1909) Echinodermata. In: Etheridge, R. & Junr, J. P. (Eds.), Scientific results of the Trawling Expedition of H. M. C. S. \" Thetis \" off the coast of New South Wales, in February and March, 1898. The Australian Museum Memoir, Sydney, pp. 519 - 564.","Ljungman, A. V. (1867) Ophiuroidea viventia huc usque cognita, enumerat. Ofversigt af Konglige Vetenskaps-Akademiens Forhandlingar, 23, 303 - 336.","Matsumoto, H. (1915) A new classification of the Ophiuroidea: with descriptions of new genera and species. Proceedings of the Academy of Natural Sciences of Philadelphia, 67, 43 - 92.","Matsumoto, H. (1917) A monograph of japanese Ophiuroidea, arranged according to a new classification. Journal of the College Science, Imperial University Tokyo, 38, 1 - 408.","Hertz, M. (1927) Die Ophiuroiden der Deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse der Deutchen Tiefsee- Expedition auf den Damfer Valdivia 1898 - 1899, 22, 59 - 122.","Thomas, L. P. (1962) The shallow water amphiurid brittle stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of Marine Science, 12, 623 - 694.","Madsen, F. J. (1970) West African Ophiuroids. Atlantide Report, 11, 151 - 243.","Paterson, G. L. J. (1985) The deep-sea Ophiuroidea of the north Atlantic Ocean. Bulletin of the British Museum (Natural History) Zoology, 49, 1 - 162.","Ljungman, A. V. (1865) Tillagg till kannedom af Skandinaviens Ophiurider. Ofversigt af Kgl. Vetenskaps-Akademiens Forhandlingar 1863, 27, 359 - 367.","Forbes, E. (1843) On the Radiata of the Eastern Mediterranean. Part I., Ophiuridae. Transactions of the Linnean society of London, 19, 143 - 153. https: // doi. org / 10.1111 / j. 1096 - 3642.1842. tb 00360. x","Hutton, F. W. (1878) Notes on some New Zealand Echinodermata, with description of new species. Transactions of the New Zealand Institute, 9, 305 - 308.","Lyman, T. (1865) Ophiuridae and Astrophytidae. University Press: Welch, Bigelow, & Company, Cambridge, 203 pp.","Rathbun, R. (1879) A List of the Brazilian Echinoderms: With Notes on Their Distribution, Etc. Transactions of the Connecticut Academy of Arts and Science, New Haven, 40 pp. https: // doi. org / 10.5962 / bhl. title. 16126","Thomas, L. P. (1966) A revision of the tropical American species of Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16, 827 - 833.","Tommasi, L. R. (1970) Os ofiuroides recentes do Brasil e de regioes vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 20, 1 - 146.","Doderlein, L. (1910) Asteroidea, Ophiuroidea, Echinoidea. In: Schultze, L. (Ed.) Zoologische und Anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Sudafrika ausgefuhrt 1903 - 1905 (Jenaer Denkschrift 16). Verlag Von Gustav Fischer, Jena, pp. 246 - 258.","Ayres, W. O. (1852) An account of the structure of the Ophiuridae and a description and drawings of a new species belonging to the genus Ophiolepis. Proceedings Boston Society Natural History, 4, 133 - 135.","Leach, W. E., Nodder, F. P., Nodder, R. P. & George, S. (1815) The zoological miscellany: being descriptions of new, or interesting animals. B. McMillan for E. Nodder & Son and sold by all booksellers, London, 144 pp. https: // doi. org / 10.5962 / bhl. title. 41372"]}

Symbiotic associations with subcuticular bacteria (SCB) have been identified and studied in many echinoderms, including the SCB of the brooding brittle star, Amphipholis squamata. Previous studies on the SCB of A. squamata placed the isolated bacterium, designated as AS1, in the genus Vibrio (Gammaproteobacteria), but subsequent studies suggested that the SCB of echinoderms belong to the Alphaproteobacteria. This study examines the taxonomic composition of SCB associated with A. squamata from the Northwest Atlantic using the 16S rRNA gene and next generation sequencing. Results show the presence of a single dominant bacterial type, within the Roseobacter clade, family Rhodobacteraceae, which composes 70%-80% of the A. squamata microbiome. These Rhodobacteraceae sequences were identified as members of the genus Octadecabacter. Additionally, the original isolate, AS1, from the brittle star A. squamata also belongs in the genus Octadecabacter based on Sanger sequencing of cloned 16S rRNA gene sequences. By comparison, adjacent seawater and sediment porewater communities were significantly more diverse, hosting bacteria in the phyla Proteobacteria, Bacteroidetes, Cyanobacteria, Verrucomicrobia and Actinobacteria. Thus, a distinct SCB community is present in A. squamata that is dominated by a member of the genus Octadecabacter and is identical to the original isolate, AS1, from this brittle star.

Brooding in ophiuroids (Echinodermata: Ophiuroidea) is a highly-developed reproduction strategy and has evolved independently several times. In South African waters, brooding occurs both in the very small amphiurid Amphipholis squamata, which is hermaphroditic, and in the large ophiodermatid Ophioderma wahlbergii, which is gonochoric. For the first time, microcomputed tomography was applied to visualise and compare brooding and brooded young of these two brittle star species from two distantly related families. The scanned A. squamata brooded six young, whereas O. wahlbergii brooded 13 young. Volume measurements of the adults revealed that A. squamata dedicated about 25% of its disc volume to brooding and O. wahlbergii about 14%. Although the brooded young of O. wahlbergii are thought to be among the largest of all known ophiuroids, young of A. squamata were four times larger in relation to adult body disc volume. Other than differences in the sizes of brooded young, these distantly related species have e...

Life history traits are among the major forces influencing the spatial organisation of biodiversity. Brooding species, lacking a planktonic larval phase, have a weak potential for dis - persal and are prone to displaying strong spatial genetic structures of their populations. Selfreproduction allows a single individual to establish a new population. Using nuclear markers to assign specimens to species (1004 specimens) and sequences of the 16S mitochondrial gene (for a subset of 479 specimens) to estimate genetic differentiation, we analyzed spatial and bathymetric sampling of 14 locations along the French Mediterranean coast in order to investigate the genetic effects of brooding and self-reproduction on the Amphipholis squamata species complex. The spatial organisation of the complex appeared chaotic, illustrating the random nature of dispersal in these brooding organisms. Bayesian dating confirmed the old age of the species complex (approximately 10 million yr). The different species displayed contrasted levels of genetic diversity and differentiation, despite their similar and extreme self-reproduction rates. This study illustrates the role of stochastic dispersal on species assemblages and genetic structure, and suggests a strong influence of past demographic history on population genetic structure of co-distributed species.

The limited availability of test organisms that represent tropical and deeper water environments is a significant concern when assessing the risk of contaminants in these environments. Amphipholis squamata (Delle Chiaje 1828) is a widely distributed brittle star with many phylogenetic clades reported from different latitudes, and it also occurs from the intertidal zone to a depth of ∼1300 m. In the present study, the effect of copper on four behavioural responses and mortality of A. squamata were quantified at four different temperatures including 25, 20, 15 and 10 °C. At 25 °C the four behavioural responses and mortality were relatively sensitive to copper, with 96 h EC50 values of 25 (confidence interval 18–44), 24 (7–26), 32 (24–41), 29 (9–41) μg L−1 for the measured ability to turn from the oral surface up to oral surface down, curling behaviour, tube foot movement, and tube foot retraction respectively. The average 96-h LC50 value for copper at 25 °C was 46 μg L−1. Some endpoints investigated showed significant effects of reduced temperature compared to the optimal temperature. These effects were enhanced with increasing copper concentrations and significant differences in copper toxicity between temperature treatments were most notable when measuring the ability to turn from the oral surface up to oral surface down where the EC50 changed from 25 (18 to 44) to 6 (−18 to 14) μg L−1 with a reduction of temperature from 25 to 15 °C. The results showed that A. squamata is relatively sensitive to copper and that further investigation into the effects of other stressors on these endpoints is warranted.

Amphipholis squamata (DELLE CHIAJE, 1828) complex (UF- 6449, KU 594365) Material. 1 spm, St. 5; 27 spms, St. 8; 20 spms, St. 9 & 7 spms, St. 10 Remarks. Amphipholis is currently composed of 26 recognized species. A. squamata is a tiny organism (DDA. squamata was revealed to be a species complex comprising numerous cryptic lineages (Sponer 2002; Boissin et al. 2008 a). Specimens from R��union are cream coloured, sometimes with a pale brown disc, and belong to the tropical lineage E (Boissin 2008). They were collected from under rocks and among coral rubble, commonly along the arms of Ophiocoma species. This association has also been noticed at Mer Island (Torres Strait), Hawaii and French Polynesia (Devaney 1974). Abundance: 3. Distribution. A cosmopolitan species complex known from all oceans except polar ones, from shore to 1300 m (Gage et al. 1983). The species was already recorded from Rodrigues (Rowe & Richmond 2004) and R��union (St��hr et al. 2008)., Published as part of Boissin, Emilie, Hoareau, Thierry B., Paulay, Gustav & Bruggemann, J. Henrich, 2016, Shallow-water reef ophiuroids (Echinodermata: Ophiuroidea) of R��union (Mascarene Islands), with biogeographic considerations, pp. 273-297 in Zootaxa 4098 (2) on pages 277-278, DOI: 10.11646/zootaxa.4098.2.4, http://zenodo.org/record/255654, {"references":["Devaney, D. M. (1974) Shallow water Asterozoans of Southeastern Polynesia. II. Ophiuroidea. Micronesica, 10 (1), 105 - 204.","Gage, J. D., Pearson, M., Clark, A. M., Paterson, G. L. J. & Tyler, P. A. (1983) Echinoderms of the Rockall Trough and adjacent areas. I. Crinoidea, Asteroidea and Ophiuroidea. Bulletin of the British Museum (Natural History), Zoology, 45, 263 - 308. http: // dx. doi. org / 10.5962 / bhl. part. 28002","Rowe, F. W. E. & Richmond, M. D. (2004) A preliminary account of the shallow-water echinoderms of Rodrigues, Mauritius, western Indian Ocean. Journal of Natural History, 3273 - 3314. http: // dx. doi. org / 10.1080 / 002229301695105","Stohr, S., Conand, C. & Boissin, E. (2008) Brittle stars (Echinodermata: Ophiuroidea) from La Reunion and the systematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnean Society, 153, 545 - 560. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2008.00401. x"]}

The species of Ophiuroidea that commonly occur on coral reefs can be found on different substrates and associated with other organisms including algae, sponges, and corals. This study evaluated the temporal variability of the ophiuroid fauna associated with the alga Amphiroa fragilissima on the Ponta Verde coral reef on the coast of Maceio in Alagoas, Northeastern Brazil. Replicate samples of algal substrate were collected at low tide during in the dry and rainy seasons, over a 10-year period, from 1998–2007. Eight species of Ophiuroidea were identified, with a total of 8736 individuals associated with A. fragilissima. The most dominant and frequent species of Ophiuroidea were Amphipholis squamata and Ophiactis savignyi; all other species were rare. Cluster analysis and nMDS detected four groups, with two groups each occurring only in summer or winter periods, as well as other groups occurring in both periods.

The black brittle star Ophiocomina nigra, common in the English Channel, is known to produce mucus when attacked. This mucus, already known for its antifouling capabilities and its role in the feeding and the locomotion behaviours of the brittle star, also emits weak light. We describe and characterize this emission of bioluminescence, thanks to a chemical triggering by hydrogen peroxide. It appears that the light emitted is 1000 times less intense than the light emitted by other brittle star species (Ophiopsila aranea and Amphipholis squamata). The luminous capabilities are homogeneously spread along the arms of the brittle star, what goes against the use of bioluminescence as a sacrificial lure. The mechanical stimu­lation of arms before chemical triggering strongly enhances the luminous capabilities of the brittle star. Luminous mucus emission can be associated with other defensive function, such as a smoke screen effect or a burglar alarm, but these two functions require intense light emissions. The fact that the luminous component is excreted outside the body might be in favour of the aposematic use of light, i.e., as a signal to warn predators of the toxicity or unpalatability of its prey.

International audience; This study aims to identify the composition and the distribution of macrobenthic communities in the Rade de Cherbourg (RdC), on the north coast of the Cotentin Peninsula (Normandy, France). The results show the presence of six main benthic communities: (1) Crepidula fornicata banks in coarse mixed sediment, (2) Amphipholis squamata and Apseudopsis latreillii in mixed sediment, (3) Melinna palmata in muddy sand, (4) Melinna palmata in mixed muddy fine sand, (5) Spio decoratus in fine sand and (6) Spio decoratus and Apseudopsis latreillii in very fine and fine sand (from EUNIS classification). The RdC appears as a rich soft bottom macrobenthic habitat in a semi-enclosed bay, surrounded by hard-bottom environments. However, current and future projects related to human activities (harbour extension, salmon farming) could impact the benthic communities. Moreover, the structure and composition of the muddy fine sand and fine sand communities, which are dominant within the area of the RdC, show similarities with comparable communities found in the western English Channel under the influence of Atlantic waters.

Within the brittle star genus Amphipholis, individuals are generally small, gonochoric, broadcast spawners, and species often have restricted distribution areas. In contrast, A. squamata (Ophiuroidea, Echinodermata)—a species complex found worldwide—is hermaphroditic and broods its young. We used 2 complementary methods to estimate the selfing rates of this species in the north- western Mediterranean Sea. First, we analysed 190 specimens from Les Goudes (France) using 3 co- dominant microsatellites and deduced the selfing rates from heterozygote deficiency, developing an original method to infer confidence intervals. Not a single heterozygote was discovered, leading to estimated selfing rates of 1.0 for all species, with an inferior limit between 0.77 and 0.98. In 7 other European localities, not a single heterozygote was found. Second, we analysed 91 brooders and their 369 offspring using 7 nuclear markers. All progeny had DNA fingerprints identical to those of the par- ents, confirming that selfing rates are very high. Such high selfing rates, by contributing to coloniza- tion success (Baker's rule), may solve the paradox of cosmopolitism for these ophiuroids that lack a larval phase that would, in theory, ensure dispersal.

Carranza, A., Borges, M., Rodriguez, M. & Borthagaray, A.I. Ophiuroidea (Echinodermata) from La Coronilla-Cerro Verde (Uruguay): a new record for the Uruguayan coast. Biota Neotrop. Sep/Dez 2007 vol. 7, no. 3 http://www.biotaneotropica.org.br/v7n3/pt/abstract?article+bn01607032007. ISSN 1676-0603. La Coronilla-Cerro verde has been proposed as the first marine protected area in Uruguay. As part of a detailed benthic biodiversity assessment at the reserve, we analyzed the ophiuroid fauna collected in the intertidal and shallow subtidal during 2005 and 2006. Three species of ophiuroids were identified: Amphioplus lucyae, Amphipholis squamata and Amphiodia sp. Only two species belonging to the genus Amphiodia (A. pulchella and A. planispina) have been previously mentioned for Uruguayan waters, indicating that the unidentified species found in this study constitutes a previously unreported species.

In this study, we present descriptions, illustrations, comments, and bathymetric and geographic distributions of the brittle star species related to the estuary region of Camamu Bay, located in the State of Bahia, Brazil. The brittle star fauna lives on biological substrates, sand bottoms, mud and rubble in the Camamu Bay and comprises 12 species divided into five families. Almost all of them are common in the tropical and subtropical fauna of the regions of shallow water.Ophiophragmus filograneusis reported for the first time in Bahia, and nine other species are recorded for the first time in Camamu Bay:Amphipholis januarii, Amphipholis squamata, Ophiophragmus filograneus, Ophiostigma isocanthum,Ophioderma cinerea, Ophioderma januarii, Ophiactis lymani, Ophiactis savignyi andOphiocoma echinata.The results suggest that the ophiuroid assemblages are strongly affected by marine currents as well as by different kinds of bottom substrate.

Morphological and molecular data on brittle stars from the Saint Peter and Saint Paul Archipelago, a very small isolated group of islets situated between South America and Africa, are hereby presented. We found no endemic ophiuroids. Instead, the five species, Amphipholis squamata, Ophiactis lymani, Ophiactis savignyi, Ophiocomella ophiactoides and Ophiothrix (Ophiothrix) angulata, are either cosmopolitan or derived from the equatorial western Atlantic region. Results indicated a western Atlantic colonization and highlighted the existence of cryptic species in the genus Ophiactis.

In this study, an overview of the current species composition of ophiuroids off Helgoland, German Bight (North Sea) is given. In addition, abundance and distribution of ophiuroids on different types of soft bottom sediments taken by van Veen grab samples around Helgoland were recorded. The literature was reviewed in order to outline the diversity of ophiuroid species over the past 130 years in the inner German Bight. In the historical literature, quantitative references often apply to verbal descriptions and thus make comparisons to modern data and assessment of possible changes in abundance rather subjective.In total, six ophiuroid species were identified off Helgoland: Acrocnida brachiata, Amphiura filiformis, Amphipholis squamata, Ophiothrix fragilis, Ophiura albida and Ophiura ophiura. The species found in this study had also been reported by previous investigators and are regarded as ‘common’ species in the German Bight. Occasional findings in this area refer to Amphiura chiajei, Ophiopholis aculeata, Ophiocten affinis and Ophiura sarsi, which are regarded here as ‘rare’ species. These findings, however, do not indicate sustainable changes in the species diversity over time.Apart from Acrocnida brachiata, a newcomer in the 1970s, findings of the common species mentioned above can be dated back to 1875. Therefore, a fairly stable composition of brittle stars is represented in the inner German Bight during the past 130 years.

The role of adrenaline in the nervous control of bioluminescence in three brittlestar species, Amphiura filiformis, Amphipholis squamata, and Ophiopsila aranea, was assessed by testing two different beta-adrenergic antagonists (propranolol and labetalol) over a wide concentration range (10(-10)-10(-3)M). We compared the effects of analogues (active vs. inactive) of the same substance (L- and D-enantiomers of propranolol). Propranolol presented both specific and nonspecific effects: (i) nonspecific effects were observed at the higher concentrations tested (10(-4) and 10(-3)M) in all three species; (ii) specific effects were detected only at the lower concentrations tested (10(-6)-10(-5)M). In A. squamata, the involvement of adrenaline in the nervous control of luminescence is supported by propranolol and labetolol specific inhibition. The neuropharmacological implications of nonspecific effects, the involvement of adrenaline and the interspecific differences in the brittlestar nervous control of bioluminescence are discussed.

We conceived a microsatellite enrichment protocol in which probes of several repeat motifs are mixed during the whole procedure. The repeats found generally differed from those of the probes and were very diverse, from mononucleotides to pentanucleotides. We tested several modifications with alternative: (i) digestion/ligation buffers; (ii) polymerases; and (iii) purification methods. The simplest methods always worked as well as classical ones or even better, resulting in probably the simplest protocol for isolating microsatellites of diverse motifs from genomes with low microsatellite frequency. The proportion of positive clones, polymorphism levels and cross-specific amplification of microsatellite loci significantly vary between species.

SUMMARYStudy of the control mechanisms of light emission in invertebrates shows the involvement of several neurotransmitters. In ophiuroids, only one species (Amphipholis squamata) has so far been characterized for luminescence control, which seems to be cholinergic, with an influence of several excitatory and inhibitory neuromodulators (amino acids, catecholamines, neuropeptides S1 and S2, purines). The aim of this work is to investigate the nature of control mechanisms of light emission in three luminous ophiuroid species, A. filiformis, O. aranea and O. californica, in order to see whether or not they share common mechanisms. Luminescence induced by general depolarisation of tissues using KCl (200 mmol l–1) shows different patterns, according to species. Only A. filiformis emits light in response to acetylcholine. In this species, the involvement of both muscarinic and nicotinic receptors is proposed, since atropine and tubocurarine (at 10–3 mol l–1) inhibited 99 % and 71 %, respectively, of the light emitted. Study of the subtypes of cholinergic receptors involved in photogenesis revealed that several subtypes of muscarinic receptors might be involved. It was also clearly shown that ophiuroids did not share a common mechanism of nervous control of luminescence in all species.

The brittle star Amphipholis squamata is paradoxical in that it lacks an obvious disper- sive phase yet has a world-wide distribution. Although individuals from distant populations are mor- phologically similar, a recent phylogenetic analysis found multiple clades separated by large genetic divergences. These clades were not phylogeographically structured and genetic divergences within populations were typically as high as those amongst populations. The recent suggestion that the sym- patric colour varieties 'orange', 'dark brown', 'beige', 'black' and 'grey' represent sibling species, led us to test whether colour variety and phylogeny were congruent. Genetic distances among sequences of the mitochondrial gene 16S rRNA from the colour varieties were surprisingly high (up to 13% uncorrected distance) and phylogenetic analyses using maximum parsimony, maximum likelihood and neighbour joining gave well supported, congruent phylogenies. However, the clades were not consistent with colour variety. When clades were constrained to make colour varieties monophyletic, tree scores were always significantly worse. We conclude from the results of this study that colour varieties do not represent distinct phylogenetic lineages. We discuss the implications of our results in the light of the possibility of clonality or self-fertilization in this species.

Amphipholis squamata (Delle Chiaje, 1828) Asteria squamata Delle Chiaje, 1828: 74. Amphiura parva Hutton, 1878: 305 [synonymised by Benham, 1909]. Amphiura squamata.���Lyman, 1879: 32.���Lyman, 1882: 136. Amphiura elegans.���Farquhar, 1898: 191���192.���Farquhar, 1907: 125. Amphipholis australiana Clark, H.L., 1909 a: 540 ���541, pl. 52 (1���3) [synonymised by Clark, H.L., 1938]. Amphipholis squamata.���Benham, 1911: 152���153.���Clark, H.L., 1921: 106���107.���Mortensen, 1924: 161���162.���Koehler, 1930: 102���103.���Mortensen, 1936: 292���293.���Clark, H.L., 1938: 243���244.���Clark, H.L., 1946: 202.���Fell, 1952: 20.���Fell, 1953: 102.���Fell, 1958: 28.���Clark, A.M., 1970: fig. 1g.���Clark, A.M. & Rowe, 1971: 80 ���81, 99, fig. 27 b, pl. 13 (5).���Clark, A.M. & Courtman-Stock, 1976: 151���152, fig. 138.���Fenwick & Horning, 1980: 439.���Paterson, 1985: 91, fig. 36. Axiognathus squamata.���McKnight, 1967 a: 311���312.���McKnight, 1967 b: 325. Material Examined. Bay of Islands. TAN0906/ 33, NIWA 54873 (2). TAN0906/ 87, NIWA 55547 (1). Marlborough Sounds. TAN 1105 / 156, NIWA 77843 (1). West Coast North Island. TAN 1105 / 88, NIWA 77842 (1). Diagnosis. Disc scaled dorsally and ventrally, radial shields contiguous. Primary plates not obvious in adult animals. Two distal oral papillae: square proximal papilla, low and wide oblong distal oral papilla. Three arm spines, 2 tentacle scales. Viviparous, bioluminescent. Description. See Mortensen (1924). Distribution. New Zealand, Australia, circumglobal in temperate and tropical waters (0���1000 m)., Published as part of Mills, V. Sadie & O'Hara, Timothy D., 2013, Ophiuroids (Echinodermata; Ophiuroidea) of biogenic habitats on the continental shelf of New Zealand, pp. 401-444 in Zootaxa 3613 (5) on pages 410-411, DOI: 10.11646/zootaxa.3613.5.1, http://zenodo.org/record/222703

Amphipholis squamata is a small ophiuroid considered to be distributed worldwide except in Polar Regions. Numerous colour varieties were reported fur this species, which is also bioluminescent. It is hermaphrodite and brood protecting; both selfing and outcrossing occur. A high genetic variability was observed among adult individuals belonging to very close (Mediterranean) local populations. Three distant populations were investigated in order to characterize morphological, physiological and genetic variability. In the population of Normandy (France), Amphipholis were collected under stones of tide pools; ophiuroids from Sicily (Italy) were collected in a Cymodocea nodosa meadow from a lagoon completely isolated from the sea. since 10 years; individuals from Santa Barbara (USA) were sampled in the aquarium system of the Marine Institute. Four colour varieties are studied throughout the sampled populations and each exhibits its own capability to produce light: (1) a spotted variety is present in the 3 populations and produces light of high intensity; (2) the orange, grey and black varieties are only present in the population of Normandy and produce light of lower intensity. Genetic variations were revealed by RAPDs. Preliminary results indicate that generic structure is homogenous for each colour variety within a population while, as expected, it shows inter-population variations for the same variety. This suggests that polychromatism and bioluminescence might be good indicators of variability of genotypes only at an intra-population level.

We investigated the effects of cyclic nucleotides (cGMP, cAMP) and the phosphoinositide IP(3) on the luminescence of the ophiuroid Amphipholis squamata. The cGMP analogue, dibutyryl-cGMP, and the guanylate cyclase activator, sodium nitroprusside, had no effect on the luminescence. The cAMP analogue, dibutyryl-cAMP, and the adenylate cyclase activator, forskolin, triggered luminescence. Moreover, the adenylate cyclase inhibitor, MDL-12330A, significantly reduced ACh-induced luminescence. The phospholipase C inhibitor, U-73122, also significantly reduced ACh-induced luminescence. The results suggest that ACh-induced luminescence is mediated by both cAMP and IP(3) pathways but not by cGMP. The effects of calcium-free ASW confirmed this hypothesis. A hypothetical scheme of the transduction mechanisms involved in the intracellular control of luminescence is presented.

Individuals of Amphipholis squamata from two colour varieties (beige and black) that differ strongly in luminescence performance, were collected from the field and raised in aquaria through three subsequent generations. The ophiuroid is a brooding species and a simultaneous hermaphrodite. Progeny from mature isolated adults and from pairs of adults of the same or of different colour varieties were scored for their characters of body colour and luminescence. Both characters change with age of the individual and reach maximal expression in the adult, the body colour being more marked and the luminescence production more intense and faster. Nevertheless, juveniles of the beige and the black variety were already clearly distinct in body colour and luminescence performance. Also, although body colour became diminished over the three generations in captivity, all progeny, from juvenile to adult, displayed the same characters of body colour and luminescence as the brooding parent. This was confirmed by observation from field-collected individuals whose brooded juveniles were always of the variety of the brooding parent. During rearing, no intermediate variety occurred that would indicate cross-fertilization between varieties. Thus, individuals could have reproduced by cross-fertilization when together with another individual if only of the same variety, or by self-fertilization when alone. Therefore, our results imply that body colour and luminescence characters are genetically determined, and that the beige and the black varieties are distinct sibling species, genetically different and isolated for reproduction.

Six colour varieties were recognized in an intertidal population ofAmphipholis squamatafrom Normandy (France). Each variety exhibits its own capacity to produce light, the light emitted by the most luminous variety being 500 times more intense than that emitted by the least luminous one. Differences in luminescent capabilities observed between varieties do not seem to be due to differences in pigmentation or to be of exogenous origin. It was shown that the capability to produce light changes according to whether individuals are brooding or not, brooding individuals emitting a more intense light than non-brooding ones. This supports the defensive use of luminescence generally associated to ophiuroid production of light. However, light capabilities differ so much betweenA. squamatacolour varieties that bioluminescence could also be associated with another role in the species.

The recent isolation and characterization of the SALMFamide neuropeptides S1 (GFNSALMFamide) and S2 (SGPYSFNSGLTFamide) from the sea stars Asterias rubens aid Asterias forbesi have initiated numerous studies on their morphological localization and distribution within the phylum Echinodermata. It has been shown by immunocytochemistry and radioimmunoassay that these peptides are widely distributed in the nervous system of some asteroids, echinoids and ophiuroids. A physiological approach has also shown that S1 and S2 potentiate the luminescence of the small ophiuroid Amphipholis squamata. In the present study, S1- and S2-like immunoreactivity have been localized in A. squamata by immunocytochemistry on both wholemount preparations and histological sections. The results reveal a widespread neuronal distribution of S1-like immunoreactivity in the circumoral ring, radial nerve cord, and tube feet. S1-like immunoreactivity was found to be associated with axons and cell bodies in both the ectoneural and hyponeural components of the nervous system. S2-like immunoreactivity was detected only in the ectoneural plexus of the circumoral ring and radial nerve cord.

Amphipholis squamata is a small bioluminescent ophiuroid whose arms are the only body part to produce light. The morphology of the arms was described paying particular attention to the spinal ganglia, viz the areas of most intense luminescence. Spinal ganglia consist of five different cell types (A-E) which were studied at different stages of the photogenous reaction. Type D cells have numerous irregularlyshaped vacuoles, widespread Golgi apparatus and well-developed rough endoplasmic reticulum (RER) that show obvious ultrastructural changes after luminescence. Type D cells appear, therefore, to be the best photocyte candidate. Type B and C cells were frequently observed in the nervous system outside spinal ganglia. Type A and E cells have not been described before. Type A cells are ciliated cells and type E cells extend long processes which are intimately associated with type D cells and epidermal ciliated cells. Both type A and type E cells could take part to the stimulation pathway that triggers luminescence.

Potassium chloride and cholinergic drugs were rested on isolated arms of Amphipholis squamata; potassium chloride triggered a monophasic light emission; acetylcholine (10(-3) M) induced a luminescent response characterized by multiple flashes. ACh luminescence was increased in the presence of the cholinesterase inhibitor, eserine. Nicotinic antagonists, tubocurarine and hexamethonium chloride, had no inhibitory effect on the luminescence; muscarinic antagonists, atropine and benzotropine mesylate inhibited Amphipholis luminescence. These results suggest that the luminescence in A. squamata is mediated through a muscarinic cholinergic system. Copyright (C) 1996 Elsevier Science Inc.

Echinoderms from the shelf seas around the British Isles were examined by epifluorescence and transmission electron microscopy for the presence of sub-cuticular bacteria (SCB). Information was obtained on SCB distribution in 63 of the 88 species known from this area, and 40 of these contained SCB. The SCB were present in all five classes. Only 39% of the holothurian species had SCB. In each of the other four classes, ≥60% of the species were associated with SCB. No correlation was noted between host ecology and SCB distribution or morphology. SCB distribution appeared to be related to host phylogeny. Congeneric echinoderms almost always had or lacked SCB, and this was usually also true for co-familial species. SCB showed limited morphological diversity, with all of them fitting into a classification of three general types. There was no morphological evidence suggesting that they are chemoautotrophs or methylotrophs. Some species had two morphological types of symbiont. SCB load was quantified for four species, three of which had symbiont densities >109 SCB g-1 ash-free dry wt. SCB are probably heterotrophic and may be co-evolving with their hosts.

The echinoderm species richness of the Aerolito de Paraiso anchialine cave, on Cozumel Island, in the Mexican Caribbean, is assessed on the basis of morphological and DNA barcoding data. We included specimens from this cave system and from different open sea areas, and employed two different approaches for species delineation based on DNA barcoding data: a 2% cox1 divergence and the general mixed Yule-coalescent (GMYC) approaches. We subsequently compared the results derived from these approaches with our morphospecies discrimination. A total of 188 cox1 sequences belonging to specimens of four echinoderm classes were examined. The 2% cox1 divergence and GMYC approaches recovered 78 and 70 putative species, respectively, 24 and 22 of which corresponded to specimens from the anchialine system. Of 26 echinoderm species identified in the cave system, seven appear to be endemic to it. Among these are Copidaster carvenicola Solis-Marin & Laguarda-Figueras, 2010, two morphologically distinctive, undescribed species belonging to Asterinides and Ophionereis and four probably cryptic undescribed species originally assigned to Amphipholis squamata (Delle Chiaje, 1839), Astropecten duplicatus Gray, 1840, Copidaster lymani (AH Clark, 1948) and Ophiothrix angulata (Say, 1825). Further research and protection of this particularly fragile ecosystem becomes urgent because construction of tourism developments is planned nearby.

Arm tissues from species of brittlestars from the British Isles and North America were examined by light and electron microscopy for the presence of sub-cuticular bacteria (SCB) which were recorded in 17 of the 19 species studied. Three major groups of SCB could be distinguished on the basis of their morphology. The bacterial morphology was not correlated with the host ecology but did appear to be related to the phylogeny of the hosts. Ophiuroids of the same genera had morphologically similar SCB and this was also generally true of species from the same family. This indicates the possibility of co-evolution between the SCB and their hosts. Estimates of SCB load were made for 9 of the species by direct counting and by quantifying the amount of bacterial endotoxin present in the host tissues. SCB numbers varied interspecifically from 3.7×108 to 4.6×109 SCB per gram of ash-free brittlestar arm tissue. Endotoxin values ranged from 5.97 to 285 μg g−1 ash-free arm tissue. Calculations suggest that SCB form at least 0.1 to 1% of the total biomass of the arm tissues. There was considerable intraspecific variation in SCB load. The possible role and significance of SCB is discussed.

Adult orthonectids develop from germinal cells within a cytoplasmic matrix called a plasmodium. This is generally assumed to be formed by the parasite. In the case of Rhopalura ophiocomae, which lives in the brittle star Amphipholis squamata, the plasmodia occupying the perivisceral coelom are closely associated with the walls of the genital bursae or the gut, and they are covered by peritoneum. They have been reported to contain scattered small nuclei distinct from those within germinal cells, embryos, and adults, but the results of the present study indicate that such nuclei probably do not exist. Furthermore, electron micrographs show that some plasmodia are in continuity with the cytoplasm of contractile cells that lie beneath the peritoneum of a genital bursa or the gut of the host. The matrix of a plasmodium of R. ophiocomaeappears, therefore, to consist of cytoplasm of a contractile cell. It is proposed that after a contractile cell has been entered by an infective cell of the parasite, it hypertrophies, bulging progressively farther into the perivisceral coelom and lifting up the peritoneum, which remains in intimate contact with it.

Deviation from pentamerism is a rare phenomenon in 5-armed echinoderm species. Qualitative and quantitative data from field observations of 14 populations around the world were used for a biogeographical study of this phenomenon. Four abnormal forms presenting deviation from the classical pentamerism of echinoderms were observed within the brittle-star species Amphipholis squamata (Echinodermata: Ophiuroidea). Although these forms were rare in global populations (0·29%), they can reach important frequencies in some locations. Our results suggest that habitat structure has an influence on the occurence of abnormal forms. Moreover, deviations from pentamerism are not a heritable character but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms.

Amphipholis squamata (Delle Chiaje, 1828) (Fig. 8) Material examined. Mercator MV, TTR 15, stn AT 568, 1 ind. (DBUA 001080.07); stn AT 575, 10 inds (DBUA 001080.01), stn AT 576, 5 inds. (DBUA 001080.02), stn AT 577, 2 inds. (DBUA 001080.03); MSM01-03, stn 241, 18 inds. (DBUA 001081.01), stn 242, 9 inds. (DBUA 001081.02), stn 287, 1 ind.; Fiuza MV, TTR 14, stn AT 566, 2 inds. (DBUA 001079.01); Kidd MV, TTR 14, stn AT 528, 2 inds.; stn AT 559, 2 ind. (DBUA 001079.02), stn AT 560, 9 inds. (DBUA 001079.03), stn AT 561, 3 inds. (DBUA 001079.04); TTR MV, TTR 12, stn AT 416, 16 inds. (DBUA 001082.01); Mekn��s MV, TTR 14, stn AT 541, 1 ind.; TTR 15, stn AT 581, 1 ind. (DBUA 001080.04), stn AT 585, 1 ind. (DBUA 001080.05), stn AT 587, 1 ind. (DBUA 001080.06); MSM01-03, stn 321, 7 inds. (DBUA 001081.03), stn 335, 15 inds. (DBUA 001081.04); Yuma MV, TTR 16, stn AT 604, 4 inds. (DBUA 001083.01); Ginsburg MV, TTR 16, stn AT 607, 3 inds. (DBUA 001083.02); Jesus Baraza MV, TTR 12, stn AT 391, 1 ind. (DBUA 001082.02); Darwin MV, TTR 16, stn AT 608, 8 inds. (DBUA 001083.03); Captain Arutyunov MV, TTR 12, stn AT 399, 2 ind. (DBUA 001082.03); MSM01-03, stn 212, 1 ind. (DBUA 001081.05), stn 194, 2 inds. (DBUA 001081.06), stn 225, 4 inds. (DBUA 001081.07), stn 274, 4 inds. (DBUA 001081.08), stn 344, 2 ind. (DBUA 001081.09); Formosa Ridge, TTR 12, stn AT 388, 2 ind. (DBUA 001082.04), stn AT 389, 5 inds. (DBUA 001082.05); West of Gilbratar Strait, TTR 14, stn AT 550, 3 inds. (DBUA 001079.05), stn AT 552, 24 inds. (DBUA 001079.06); Pen Duick Escarpment, TTR 12, stn AT 406, 18 inds. (DBUA 001082.06); TTR 14, stn AT 565, 1 ind. (DBUA 001079.07); TTR 16, stn AT 600, 8 inds. (DBUA 001083.04), stn AT 602, 4 inds. (DBUA 001083.05). Ecology and distribution. Amphipholis squamata is known from several substrata (under stones, amongst algal and bryozoan turfs, rockpool weeds and occasionally on sandy bottoms) from the intertidal to 1200 m. Current distribution records suggests that it is abundant in the Eastern Atlantic from Iceland along the European margin, Mediterranean Sea, West coast of Africa, West and East coast of South Africa and Madagascar (Alva & Vadon 1989). The taxonomy of this species is currently being investigated (S. St��hr, per comm.). It is likely that this presumed cosmopolitan species will turn out to be a complex of cryptic species and that there will need to be a revision of its geographic distribution. This species can switch from deposit feeding by collecting particles within its tube feet to suspension feeding via trapping detritus in mucus. In the Gulf of Cadiz it was the most common ophiuroid, recorded from 10 mud volcanoes and at three other sites in association to carbonate chimneys and crusts as well as cold-water corals (Fig. 7 D). Young, intermediate and adult stages were found together at the same localities. This species was found at depths from 350 to 1379 m which is its deepest known record., Published as part of Rodrigues, Clara F., Paterson, Gordon L. J., Cabrinovic, Andrew & Cunha, Marina R., 2011, Deep-sea ophiuroids (Echinodermata: Ophiuroidea: Ophiurida) from the Gulf of Cadiz (NE Atlantic), pp. 1-26 in Zootaxa 2754 on pages 16-17, DOI: 10.5281/zenodo.276727, {"references":["Delle Chiaje, S. (1828) Memoire sulla storia e notomia degli animali senza vertebre del regno di Napoli. Napoli: Societa Tipographica, 3, 1 - 232.","Alva, V. & Vadon, C. (1989) Ophiuroids from the western coast of Africa (Namibia and Guinea-Bissau). Scientia Marina, 53 (4), 827 - 845."]}

1. 1. Transport rates of dissolved free amino acids for two sympatric ophiuroids were determined to assess differences due to size or life-history traits. 2. 2. Both species of brittle stars demonstrated a net flux of dissolved free amino acids from seawater, while the brooding brittle star Amphipholis squamata had significantly higher rates, greater than twice those of Ophiopholis aculeata. 3. 3. Measured rates of uptake for either species of brittle star contribute less than 5% of the respiratory substrate needed to support aerobic metabolism. 4. 4. For A. squamata, incorporation of dissolved free amino acids into protein appeared to be primarily by its symbiotic subcuticular bacteria. 5. 5. These results suggest that both uptake of amino acids and their subsequent incorporation into bacterial protein are coupled. 6. 6. However, calculations of the maximum potential uptake of dissolved amino acids by the symbiotic bacteria can account for no more than 10% of the total amino acid uptake in A. squamata. 7. 7. At the amino acid concentrations used in this study the maximum uptake of amino acids by bacteria should have been achieved.

Numerous marine species are able to assess predation risk through chemical signals, and this capacity is usually found among animals that experience intense predation and possess chemoreception ability, such as brittle stars. We investigated the occurrence of chemical alarm signals and responses in four species of brittle stars: Amphipholis squamata, Ophionereis reticulata, Ophiactis savignyi and Ophiothrix angulata. Additionally, the effect of microhabitat on alarm-signal recognition was tested for O. savignyi. Brittle-star homogenate was released above individuals in an aquarium, and the duration of their immediate escape response was noted and compared to the control. The broadest recognition was observed for A. squamata, which showed an escape reaction to the damage-released stimuli of all brittle stars tested. Similarly broad recognition was observed for individuals of O. savignyi that were collected from the same algal species occupied by A. squamata, a microhabitat where these individuals co-occurred with each other and with juveniles of O. angulata. The similar size and habit of these species probably expose them to the same predators, and therefore, the alarm signal of one species represents a real risk of predation to the others. In contrast, individuals of O. angulata and O. savignyi collected from sponges did not respond to stimuli from either conspecific or heterospecific individuals. The reduction in predation pressure granted by their chemically protected hosts seems to be responsible for this lack of response. The fourth species, O. reticulata, only responded conspecifically. This brittle star is found buried among rubble, and is not observed associated with other organisms; it is known for its cannibalistic and predatory habit.