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Health and resilience of the coral holobiont depend on diverse bacterial communities often dominated by key marine symbionts of the Endozoicomonadaceae family. The factors controlling their distribution and their functional diversity remain, however, poorly known. Here, we study the ecology of Endozoicomonadaceae at an ocean basin-scale by sampling specimens from three coral genera ( Pocillopora , Porites , Millepora ) on 99 reefs from 32 islands across the Pacific Ocean. The analysis of 2447 metabarcoding and 270 metagenomic samples reveals that each coral genus harbored a distinct new species of Endozoicomonadaceae . These species are composed of nine lineages that have distinct biogeographic patterns. The most common one, found in Pocillopora , appears to be a globally distributed symbiont with distinct metabolic capabilities, including the synthesis of amino acids and vitamins not produced by the host. The other lineages are structured partly by the host genetic lineage in Pocillopora and mainly by the geographic location in Porites . Millepora is more rarely associated to Endozoicomonadaceae . Our results show that different coral genera exhibit distinct strategies of host- Endozoicomonadaceae associations that are defined at the bacteria lineage level. published

Background: Over the last decade, several coral genomes have been sequenced allowing a better understanding of these symbiotic organisms threatened by climate change. Scleractinian corals are reef builders and are central to coral reef ecosystems, providing habitat to a great diversity of species. Results: In the frame of the Tara Pacific expedition, we assemble two coral genomes, Porites lobata and Pocillopora cf. effusa, with vastly improved contiguity that allows us to study the functional organization of these genomes. We annotate their gene catalog and report a relatively higher gene number than that found in other public coral genome sequences, 43,000 and 32,000 genes, respectively. This finding is explained by a high number of tandemly duplicated genes, accounting for almost a third of the predicted genes. We show that these duplicated genes originate from multiple and distinct duplication events throughout the coral lineage. They contribute to the amplification of gene families, mostly related to the immune system and disease resistance, which we suggest to be functionally linked to coral host resilience. Conclusions: At large, we show the importance of duplicated genes to inform the biology of reef-building corals and provide novel avenues to understand and screen for differences in stress resilience. † Benjamin Noel and France Denoeud have equal contribution.

Ophioderma zibrowii sp. nov. urn:lsid:zoobank.org:act: 7FB245FD-C8DF-402A-A7B1-FA7D048AFB04 Figs 11–12; Table 1 Diagnosis Usually naked radial shields. Seven arm spines. Eight oral papillae. Colour pattern greenish brown to dark green variegated on dorsal and ventral disc, arms banded, with light and dark spots along distal dorsal plate edges, oral frame and ventral arms lighter. Largest known size 21 mm dd. Brooder. Etymology The species is named after Helmut Zibrowius for his important contributions in collecting material for this study. The existence of brooding Ophioderma in the Mediterranean Sea might still be unknown without his effort. Material examined Holotype MEDITERRANEAN SEA • ♀ (brooding); southern Crete, Agios Pavlos; May 2012; A. Weber leg.; preserved in 95% ethanol; SMNH-Type-9188. Paratypes MEDITERRANEAN SEA • 1 ♀ (brooding); same data as for holotype; SMNH-Type-9189 • 7 specimens; Greece, Symi Island, near Rhodes, Agios Georgios; 10 Aug. 2005; R. Graille leg.; preserved in 95% ethanol; SMNH-Type-9190 to 9196. Other material MEDITERRANEAN SEA • 18 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, northwestern Crete, Elounda, Plaka Beach; 2 Jul. 2008; Emilie Aegea leg.; preserved in 95% ethanol; SMNH-102712, 113690 to 13697, 11370 to 113704, 113709 to 113712 • 21 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, Rhodes, Kallithea Bay; 9 Sep. 2005; H. Zibrowius leg.; preserved in 95% ethanol; SMNH-99657, 113982, 113985, 113987 to 113990, 113992 to 114003, 114005, 114006 • 41 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, Crete; May 2012; A. Weber leg.; preserved in 95% ethanol; SMNH-147070 • 1 specimen (dd = 15.4 mm), dried dissociated ossicles and ossicles on two SEM stubs; same data as for preceding; SMNH-178455. Description Holotype (Fig. 11) Disc diameter 14.3 mm, one intact arm, all others removed for molecular analysis. Dorsal disc with dense cover of granules, radial shields naked, widely separated, visible part oval (Fig. 11A). Granules and scales crowd onto basal arms. Dorsal arm plates single on all joints, flat, overlapping, broadly fanshaped, twice as wide as long, with straight edges, wider distally than proximally (Fig. 1D). Up to seven conical, slightly pointed arm spines, about equal in length, half as long as arm joint, ventralmost spine thicker than others and blunt (Fig. 7C). Ventral disc covered by granules similar to dorsal disc. Adoral shields and oral plates covered by granules, oral shields naked (Fig. 11B). Seven to eight oral papillae, Lyman’s ossicle elongated, pointing into mouth slit, adoral shield spines two, block shaped, other papillae smaller, conical (Fig. 11E). Tooth papillae larger, conical, two in line with oral papillae. Oral shields triangular, twice as wide as long, straight distal edge, obtuse rounded proximal angle. Madreporite as wide as long. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules, similar to disc granules. Proximal genital slits parallel to and as long as first and second joints. Distal genital slit parallel to and as long as fourth joint, at distance from disc edge. Bursae filled with brooded juveniles (Fig. 11F). Dorsal disc slightly variegated dark brown-green with grey, dorsal arms greyish green with brown bands, each light and dark band about three joints wide, a series of larger white and smaller black spots along the distal edge of each dorsal arm plate (Fig. 11A, D). Ventral disc as dark as dorsal, oral frame and ventral arms lighter grey-green (Fig. 11B). Arm spines grey-green with white tips, except ventralmost spine, which is white (Fig. 11 C–D). Oral papillae and tentacle scales white (Fig. 11C, E). Paratypes Disc diameter 9.7–13.5 mm.All have up to seven arm spines. Most paratypes have a green colour pattern similar to the holotype, one specimen is olive-brown. One paratype with granule-covered radial shields, all others naked. Oral papillae eight, in two specimens nine, but variable between jaws in an individual. Two specimens with two dorsal arm plates on 2–3 proximal joints on 1–3 arms. One specimen with 2–3 dorsal arm plates on proximal joints on three arms, on seven joints on one of those arms. One specimen has a parasitic snail (Ersilia Monterosato, 1872 sp.) on one arm. All paratypes lack part of an arm, which was used for molecular analysis. Non-paratypes (SMNH-147070) Disc diameter 8.4–20.5 mm. Specimens smaller than 10 mm dd had six arm spines, all others seven. Most specimens have a green colour pattern similar to the holotype, several were dark olive with numerous white granules scattered across the disc. Out of the 42 specimens, a single one had granule-covered radial shields, all others with naked radial shields. Most of the specimens above 10 mm dd had at least two proximal arm joints on at least one arm with two dorsal arm plates, up to seven joints and several arms with double plates in one animal. The plates were usually of equal size or there was a hairline crack along the midline of the plate. Rarely, a third dorsal arm plate was present on one or two joints. Oral papillae eight, rarely nine, but variable between jaws in an individual and not correlated to size. These specimens were used for thermotolerance and regeneration experiments. These specimens were kept in an aquarium for a year where experiments of thermotolerance and regeneration were conducted. Therefore, most arms show signs of regeneration. One specimen had bright white distal arms but no sign of healed breaks. Maximum size recorded for O. zibrowii sp. nov. was 21 mm dd (Weber et al. 2014). Skeletal elements (Fig. 12) Non-type specimen of 15.4 mm dd (SMNH-178455), completely dissociated. Radial shields elongated isosceles triangular, external surface with embossed oval of more finely meshed stereom in center of domed and thickened proximal half (only visible part in intact skeleton), row of eight pores curves around proximal dome (Fig. 12A, C). On inner side of radial shield, a large pore on distal half and a round patch with denser stereom, two low distal bulbs separated by a furrow (Fig. 12B). Dental plate consists of several pieces, sockets not penetrating the plate, bordered by low rim, tooth papillae sockets smaller, round, tooth sockets wide, oval (Fig. 12D). Adradial genital plate blade-like flat with bulbous distal end and large pore, distal pit with knob on one side, three ridges on other side (Fig. 12 E–F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally (Fig. 12G). Oral plates longer than high, middle part markedly lower than ends, abradial muscle fossa rounded triangular, not reaching upper edge (Fig. 12H). On adradial oral plate proximoventral part four oral papilla sockets and pores, at proximoventral edge four granule sockets and pores (Fig. 12I). Vertebrae typically zygospondylus, with large dorsal and small ventral muscle flanges, no growth rings obvious (Fig. 12 J–K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening (Fig. 12 L–M). Two spurs at proximal external edge and counterparts on internal distal edge of lateral arm plates. Internal lateral arm plate with vertical row of three pores just below the plate centre, distal to curved vertical ridge, fourth pore in tentacle opening edge, small ventral knob, protruding beyond plate edge. Spine articulations inset in the distal plate edge with one thickened lobe, separating muscle and nerve opening. Remarks The species determination rests mainly on the genetic analyses. We chose the predominant genetic group (C5) to describe a new species, but due to its short reproduction period (May–June, Weber et al. 2014) most specimens of this cluster happen to have been collected outside the brooding time. Since brooding is one of the key characters of this species, we felt that the holotype should be a brooding female. It is not ideal that the chosen animal has only a single intact arm remaining, but in a pentaradially symmetric animal one arm is sufficient and representative of the others. Morphological differentiation from O. longicauda rests on the usually naked radial shields and the single, flat arm plates. Also, O. zibrowii sp. nov. usually lacks red pigments which are common in O. longicauda, whereas green pigments are common in O. zibrowii sp. nov. but absent in O. longicauda. In addition, O. zibrowii sp. nov. commonly has a series of white and black spots along the distal edge of the dorsal arm plates, which are shared with O. hybrida sp. nov. and O. guineense. The radial shields differ microscopically from those of O. longicauda in the clear demarcation of an embossed part with finer meshed stereom, representing the naked visible part of the radial shields in the intact animal. The major part of the radial shield is overlapped and covered by disc scales, obscuring its true shape. Possibly, the more elongated oral plates and higher lateral arm plates in O. zibrowii sp. nov. distinguish the species, but these small differences need to be studied with morphometric methods and larger samples. Brooding females have been collected from Cyprus and Lebanon, but they belong to another genetic cluster (C6). Morphologically they cannot be distinguished at this time and we consider them to be allopatric populations within the same species. They are excluded from the concept of the new species O. zibrowii sp. nov., as new evidence may in the future lead to them being recognized as a different species.

Ophioderma hybrida sp. nov. urn:lsid:zoobank.org:act: 72379251-8181-4B42-8 CB 6-881F0C144526 Figs 9���10; Tables 1���2 Diagnosis Species of Ophioderma with variable morphology. Dorsal arm plates single or as several pieces. Seven to 10 equal arm spines. Eight oral papillae. Colour pattern: disc dark brown or olive dorsally and ventrally, often with white spots. Dorsal arms dark brown with light brown or white and black spots, ventrally lighter brown. Maximum known size about 19 mm dd. Etymology This species has likely evolved by an ancient hybridization event between two species (O. longicauda and O. zibrowii sp. nov. or their respective ancestors), which the Latin word hybrida alludes to. It also refers to the mixed morphology of the species. Material examined Holotype MEDITERRANEAN SEA ��� Tunisia, Kelibia; 36��50���7.31��� N, 11��7���6.20��� E; depth 0.3���1.5 m; 3 Jul. 2013; Zined Marzouk & Alexandra Weber leg.; preserved in 95% ethanol; SMNH-Type-9185. Paratypes MEDITERRANEAN SEA ��� 11 specimens; same data as for holotype; SMNH-Type-9186 ��� 8 specimens; Tunisia, Monastir; 35��46���28.83��� N, 10��50���18.93��� E; depth 2���5 m; 25 Jun. 2013; A. Weber leg.; preserved in 95% ethanol; SMNH-Type-9187. Other material MEDITERRANEAN SEA ��� 3 specimens (with dorsal disc removed, but present); same data as for holotype; SMNH-178454 ��� 3 specimens; Tunisia, Tunis; 1849; ��berg leg.; SMNH- 35614, 123374, 123375. Description Holotype (Fig. 9) Disc diameter 17.4 mm, two arms cut off near disc but present (Fig. 9 A���B). Radial shields covered by disc granules (Fig. 9A). Dorsal arm plates weakly tumid, as 1���3 pieces on about first 30 joints, when single twice as wide as long, distal edge straight or concave (Fig. 9 C���D). Ten arm spines, all about equal in length, ventralmost slightly wider. Ventral arm plates overlapping, squared, distal edge convex (Fig. 9E). Two oval scale-like tentacle scales, superimposed on the ventralmost spine. Ventral disc covered by granules similar to dorsal disc. Eight oral papillae, short, weakly pointed to round, Lyman���s ossicle large, pointing into mouth slit (Fig. 9F). Tooth papillae larger than oral papillae, teeth rounded block-shaped. Oral shield almost twice as wide as long, with wide proximal angle, distal edge slightly convex. Madreporite larger, almost as long as wide (Fig. 9B). Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm (Fig. 9F). Adoral shields covered with granules, similar to disc granules. Proximal genital slits angled from oral shield to second arm joint, distal genital slits at disc edge, two arm joints long. Disk dark brown dorsally with radiating lighter brown pattern and scattered white spots (Fig. 9A). Dorsal arms dark brown with light brown and dark spots along the distal plate edges and on some lateral edges (Fig. 9 C���D). Ventral disc and arms light greenish brown, oral shields darker (Fig. 9B). Variation The range of morphological variation is presented in Table 2 and on Fig. 10. The specimens vary in size from 9.8 to 18.7 mm disc diameter. Among the 23 examined specimens, the number of dorsal arm plate pieces varies considerably. Multiple plates or pieces have been found on the proximal arms, until about joint number 30, but not on all joints, and sometimes not on all arms (Fig. 10E). In most specimens, there are only two pieces, but in a few animals, multiple pieces were observed (Fig. 10E, H, N). Some specimens have only single dorsal arm plates (Fig. 10B, K). More pieces are present in the larger specimens but there appears to be only a weak size relation for this trait. Likewise, the number of arm spines and oral papillae is not related to disc diameter. All but two animals have granule covered radial shields. Colour variations are limited. All examined specimens have a dark brown or olive dorsal disc with varying white patterning (from spots to radiating lines) and a dark ventral disc, similar or lighter oral frame and ventral arms. The dorsal arms show alternating light and dark bands and a series of light and dark spots along the distal edge of the dorsal arm plates. Other specimens SMNH- 35614 15.5 mm dd, SMNH-123374 16.3 mm dd, SMNH-123375 14.3 mm dd. Colour faded to light brown. Most of the granules rubbed off the discs, and status of the radial shields as naked/granule covered cannot be assessed, but they have an exposed area, devoid of scales. Eight oral papillae. Two to three dorsal arm plates on single joints (SMNH-123374), up to nine joints (SMNH- 123375), up to 12 joints (SMNH-35614), variable between arms. Juveniles present in the bursae. Remarks The origin of this species through an ancient hybridization event between O. longicauda and O. zibrowii sp. nov. (or their ancestors) (Weber et al. 2019) is further supported by its high morphological variability. Multiple dorsal arm plates are typical of O. longicauda, and in some specimens of O. hybrida sp. nov. the most proximal joints bear several plates, forming a crowded mosaic pattern similar to O. longicauda. The usually granule covered radial shields are shared with all species except O. zibrowii sp. nov. (Table 1) and are probably evidence of O. longicauda (or its ancestor) being one of the ancestral species. The variation of arm spines and oral papillae overlap with both ancestral species and O. africana sp. nov., but O. guineense has a higher number of arm spines. Ophioderma hybrida sp. nov. shares the vivid colour pattern of the arms with O. zibrowii sp. nov. and O. guineense, whereas O. africana sp. nov. and O. longicauda are darker and the spots on the arms are less obvious in the former. So far, O. hybrida sp. nov. appears to be a smaller species with disc diameter under 20 mm, but more material is needed to confirm this. We believe that the reproductive mode of O. hybrida sp. nov. is brooding given that brooding individuals have been collected in Tunisia (St��hr et al. 2009) and the ones examined in this study are morphologically more similar to O. hybrida sp. nov. than to O. zibrowii sp. nov., because they have more dorsal arm plates per joint. Yet, we cannot confirm it, as the brooding specimens from Tunisia were too old for genetic analyses (1849 and 1924; PCR amplification failed). In addition, all other O. hybrida sp. nov. specimens have been collected outside the reproductive season so examination of their gonads was inconclusive in determining their reproductive strategy., Published as part of St��hr, Sabine, Weber, Alexandra Anh-Thu, Boissin, Emilie & Chenuil, Anne, 2020, Resolving the Ophioderma longicauda (Echinodermata: Ophiuroidea) cryptic species complex: five sisters, three of them new, pp. 1-37 in European Journal of Taxonomy 600 on pages 22-24, DOI: 10.5852/ejt.2020.600, http://zenodo.org/record/3637773, {"references":["Weber A. A. - T., Stohr S. & Chenuil A. 2019. Species delimitation in the presence of strong incomplete lineage sorting and hybridization: lessons from Ophioderma (Ophiuroidea: Echinodermata). Molecular Phylogenetics and Evolution 131: 138 - 148. https: // doi. org / 10.1016 / j. ympev. 2018.11.014","Stohr S., Boissin E. & Chenuil A. 2009. Potential cryptic speciation in Mediterranean populations of Ophioderma (Echinodermata: Ophiuroidea). Zootaxa 2071: 1 - 20. https: // doi. org / 10.11646 / zootaxa. 2071.1.1"]}

Ophioderma africana sp. nov. urn:lsid:zoobank.org:act: 64021157-EA56-449 B- 9BCB-FD02F10CC67F Figs 7–8; Table 1 Diagnosis Mostly single dorsal arm plates, but several plates on some joints, eight arm spines, eight oral papillae. Colour dark brown dorsally, ventral disc as dark, ventral arms lighter brown, black and white spots along distal edge of dorsal arm plates. Maximum size under 19 mm dd (26 mm dd if non-type material is considered). Etymology The species is named for its area of collection, off (West) Africa. Material examined Holotype TROPICAL ATLANTIC OCEAN • Senegal, Dakar, Gorée Island, shipwreck Tacoma; depth 10 m; 28 Aug. 2007; H. Zibrowius leg.; scuba diving, hand collected; under stones; preserved in 95% ethanol; SMNH-Type-7484. Paratypes TROPICAL ATLANTIC OCEAN • 22 specimens (6 sampled for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); same data as for holotype; SMNH-Type-7485 • 1 specimen, ossicles, 4 remaining arms, 2 SEM stubs with ossicles; same data as for holotype; dried; SMNH-Type-7486 • 11 specimens (including a separate vial with eggs that were spawned during collection); same data as for holotype; initially in formalin, then 80% ethanol; SMNH-Type-7487 • 4 specimens (2 used for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); Senegal, Dakar, Cap Manuel; depth 7 m; 6 Sep. 2007; H. Zibrowius leg.; under stones; scuba diving, hand collected; preserved in 95% ethanol; SMNH-Type-7488 • 4 specimens (2 used for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); Senegal, Madelene Islands; depth 10–12 m; 9 Sep. 2007; H. Zibrowius leg.; among piled up rocks; scuba diving, hand collecting; preserved in 95% ethanol; SMNH-Type-9184. Other material TROPICAL ATLANTIC OCEAN • 6 specimens (labelled as O. longicauda); Senegal, between Gorée and Tiaroye; depth 15–20 m; Feb.–Mar. 1952; Thorson leg.; preserved in ethanol; NHMD-225514 • 6 specimens (labelled as O. longicauda); Senegal, Dakar; Jan. 1928; Sudan exped.; H. Madsen leg.; dried; partial sample; NHMD- 225517 • 10 specimens (labelled as O. longicauda); Dakar; 6 Jan. 1928; Sudan exped.; H. Madsen & O. Hagerup leg.; preserved in in ethanol; partial sample; NHMD-225523 • 1 specimen (labelled as O. longicauda); Senegal, Dakar, rocky coast; 3 Jan. 1928; H. Madsen leg.; preserved in ethanol; NHMD-225524. Description Holotype (Fig. 7) Disc diameter 18.7 mm, disc completely covered by small granules dorsally and ventrally (Fig. 7 A–C), including the radial shields (Fig. 7B). Disc scales with granules crowding onto arm base, covering first dorsal arm plate except for a mid-distal portion, second plate also covered by scales and granules laterally (Fig. 7 D–E). Dorsal arm plates flat, 2.5 times as wide as long, with straight distal edge, contiguous, mostly single, but joints with two plates common (often unequal in size) (Fig. 7 D–E). Eight arm spines, ventralmost one thicker and slightly longer than others (Fig. 7F). Ventral disc covered by granules similar to dorsal disc. Eight conical oral papillae, Lyman’s ossicle and adoral shield spines hardly distinguishable (Fig. 7G). Two larger tooth papillae at jaw tips in line with oral papillae, blocklike teeth. Oral shields rounded triangular, slightly wider than long, madreporite slightly wider than other oral shields and with central round depression. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules, similar to disc in size. Proximal genital slits about one joint long, distal slits about three joints long (Fig. 7C). Ventral arm plates contiguous, as wide as long, with wide proximal angle, convex distal edge (Fig. 7F). Two oval scale-like tentacle scales, superimposed on the ventralmost spine. Disc dark brown dorsally, with some lighter brown patches. Dorsal arms dark brown, distally lighter and banded. Dorsal arm plates with small light brown and black dots along distal edge. Ventral disc as dark as dorsal, oral frame and ventral arms light orange brown (Fig. 7A, C). Paratypes Size range 12–18.4 mm dd. The specimens have dark brown to almost black dorsal discs (on some individuals with lighter patches), arms with dark and lighter bands. Ventral discs as dark as dorsal, oral frames and ventral arms lighter brown (Fig. 7H). Individuals with small light spots on the disc occur. Most specimens with black dots on the distal edge of the dorsal plates, but some without. Dorsal arm plates are mostly single, but most arms with several joints with two plates. Skeletal elements (Fig. 8) From specimen of 15.7 mm dd (SMNH-Type-7486) with mostly single dorsal arm plates but some small additional plates on few joints. Seven oral papillae, all similar except Lyman’s ossicle. Up to eight arm spines. Radial shields isosceles triangular with deeply concave abradial edge and abradial distal processes, distal center domed with small pores (the domed part is not covered by scales but with granules in the intact animal, all other areas are covered by scales) (Fig. 8A, C). Inner side of radial shield, on the distal articulation surfaces, two larger oval depressions and a smaller round one between these (Fig. 8B). Dental plate consists of several pieces with non-perforating sockets for 2–3 tooth papillae and single wide teeth, granule pores not obvious (Fig. 8D). Adradial genital plate blade-like flat with bulbous distal end and large pore, distal pit with knob on one side, two distal depressions on other side (Fig. 8 E–F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally (Fig. 8G). Oral plates longer than high, middle part slightly lower than ends, adradially with four oral papilla pores on ventral edge of proximal part, small granule pores at ventral edge (Fig. 8I). Adradial distal muscle fossa wide ventrally, tapering dorsalwards, shorter than plate height, articulation area slightly higher than wide (Fig. 8H). Suture line diagonally across oral plate mid-line, separated easily during preparation on several jaws. Vertebrae typically zygospondylus, with large dorsal and small ventral muscle flanges, no growth rings obvious (Fig. 8 J–K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening (Fig. 8 L–M). Two spurs at proximal external edge and counterparts on internal distal edge of lateral arm plates. Internal lateral arm plate with vertical row of three pores just below the plate centre, fourth pore in tentacle opening edge, bent vertical proximal ridge and small ventral knob, protruding beyond plate edge. Spine articulations inset in the distal plate edge with one thickened lobe, separating muscle and nerve opening. Other specimens The specimens from NHMD possess mostly single dorsal arm plates, on few joints there are more than one and some specimens have on the most proximal joints a mosaic of slightly tumid plates. Some specimens have naked radial shields. They have 7–9 oral papillae, among which the adoral shield spine and Lyman’s ossicle are sometimes obvious, differing in shape from the other papillae. They are uniformly dark brown or variegated dark, on both dorsal and ventral sides. They range in size between 19 and 26 mm dd and are assigned to O. africana sp. nov. mostly by colour pattern and exclusion of O. longicauda from off West Africa. The specimen NHMD-225524, which measures only 5.7 mm dd, has a light brown dorsal pattern with faint spots, a cream white ventral surface, and single dorsal arm plates, similar to O. guineense and is reassigned to that species. Remarks Ophioderma africana sp. nov. shares the dark colour on both sides of the disc with O. longicauda, its spine numbers fall within the variability of both O. longicauda and O. guineense, it has as few oral papillae as O. longicauda (Table 1). These characters are however variable and small specimens may be difficult to identify. Nuclear and mitochondrial data were distinct from O. longicauda and all other clusters found (Weber et al. 2019), placing O. africana sp. nov. in cluster C2 and COI lineage L5. Some of the animals started spawning during collecting, which indicates that the species is a broadcast spawner, not a brooder.

Ophioderma guineense Greeff, 1882 Figs 4���6; Table 1 Ophioderma guineense Greef, 1882: 156. Ophioderma longicauda var. guineense ��� Koehler 1914: 173. Ophioderma longicauda guineense ��� Madsen 1970: 225, fig. 43. ��� Ophioderma appressa ?��� ��� Koehler 1914: 175; Madsen 1970: 228. Diagnosis Predominantly single flat dorsal arm plates, up to 13 arm spines, nine oral papillae (counting from and including Lyman���s ossicle but excluding tooth papillae). Colouration in alcohol variable, dorsal disc brown with white spots and dark rings, arms banded dark and light brown to grey, ventral disc, arms and oral frame cream to light grey, dark and white spots along distal edge of dorsal arm plates. Disc diameter of largest paralectotype 29 mm. Material examined Lectotype (here designated) TROPICAL ATLANTIC OCEAN ��� West Africa, Gulf of Guinea, Islands Sao Tom�� and Rolas; cliffs at low tide; ZMH E8315. Paralectotypes (here designated) TROPICAL ATLANTIC OCEAN ��� 6 specimens (2 specimens at hand, 4 specimens examined on photographs); same data as for lectotype; ZMH E1686. Other material TROPICAL ATLANTIC OCEAN ��� 11 specimens (photographed specimens in separate bags labelled as #1���3, mitochondrial lineage L6, nuclear cluster C 1); Senegal, Dakar, Grand Thiouriba; depth 38 m; 6 Sep. 2007; H. Zibrowius leg.; from heavily fouled piece of fishing net; scuba diving, hand collected; preserved in 95% ethanol; SMNH- 95511 ��� 1 specimen (dd = 11.7 mm), dried arm pieces, dried ossicles, ossicles mounted on two SEM stubs; same data as for preceding; SMHN-178429 ��� 3 specimens (labelled as O. longicauda var. guineense); Equatorial Guinea, Gulf of Guinea, Annob��n; depth 18���30 m; Discovery exped. (stn 283); dried; NHMD- 225525. Description Lectotype (Fig. 4 A���D) Disc diameter 28.5 mm, radial shields widely separated, sunken in, concealed by granules. Scales and granules crowd onto proximal arms, obscuring dorsal plates laterally, granules rubbed off in some places. Dorsal arm plates entire almost on whole arm (single cracked plates on mid-arms), not tumid, three times as wide as long, contiguous (Fig. 3C). Three to 13 arm spines (from first spine bearing joint under disc to proximal free ones), flattened, slightly pointed, slightly longer than half arm-joint length, ventralmost spine longer and wider than others. Ventral arm plates wider than long, with convex distal edge, overlapping, except the proximalmost 3���4 which are separated by pores. Two oval scale-like tentacle scales, superimposed on the ventralmost spine. Ventral disc covered by granules similar to dorsal disc (Fig. 4D). Lyman���s ossicle elongated, reaching into the mouth slit, not obvious, adoral shield spines on some jaw edges block-like, on others as lateral papillae and not obvious, in total nine oral papillae per jaw edge. Two tooth papillae at jaw tips in line with oral papillae, slightly larger than oral papillae. Teeth rectangular with straight edges. Oral shields rounded triangular with obtuse proximal angle and straight distal edge, slightly wider than long. Madreporite larger than other oral shields, depressed. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules, similar to disc in size. Colour pattern with tiny dark spots on the dark grey-brown dorsal disc, dark and white spots on the distal edge of the dorsal arm plates, cream ventral side (Fig. 4 A���C). Paralectotypes (Fig. 4 E���H) Disc diameter 26 and 29 mm, both with up to 13 arm spines, 9���10 oral papillae. Radial shields concealed by granules, but in parts of the disc, the granules have been rubbed off (Fig. 4E, G). Proximal dorsal arm plates entire. In the largest syntype, colour faded light brown dorsally, cream with some dark brown ventrally, arms with faint dark bands. There are irregular dark rings on the pale brown dorsal disc of the smallest syntype examined (Fig. 4G). In these specimens, there are dark brown and white spots along the distal edge of the dorsal arm plates. Among the paralectotypes examined only from photographs, one is dark brown on dorsal disc and arms, and the dark colour extends onto the ventral disc, fading towards the oral frame. The other specimens have a faded brown dorsal colour and light ventral discs and arms. Some of these show faint spots on distal edges of the dorsal arm plates. The amplification of nuclear markers from the syntypes was not successful, most likely due to the long storage time and unknown initial preservation method of the specimens, which might have caused the DNA to deteriorate. Specimen from Senegal (SMNH-95511#1, Fig. 5 A���D) Disc diameter 15 mm, up to nine arm spines proximally, dorsal arm plates flat, entire along whole arm, radial shields covered by granules. Disc scales with granules crowding onto arm base, covering first dorsal arm plate except for a mid-distal portion, second plate also covered by scales and granules laterally. Dorsal arm plates overlapping, twice as wide as long, rounded fan-shaped, distal edge wider than proximal, not tumid. Ventral disc covered by similar granules as dorsal disc. Jaws with nine lateral papillae. Two larger tooth papillae at jaw tips in line with oral papillae, rectangular teeth with straight edges. Oral shields triangular, almost as long as wide. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules similar in size to disc granules. Proximal genital slits small, extending from oral shield to proximal edge of second ventral arm plate. Distal genital slits at disc edge, next to fifth arm joint. Disc dark brown dorsally, large white patches in centre. All over with small irregular white dots with black margins (ring-like). Disc light green ventrally. Oral shields with darker green centres. Arms dorsally dark brown, with lighter and darker bands, ventrally cream white with faint darker bands. Dorsal arm plates with small black spots along distal edge. Variation in other Senegalese specimens Size range 8���13 mm dd. The specimens vary in colour pattern. One specimen has large white patches over the area around the radial shields, on an otherwise dark disc. Some animals have a light brown disc with minute, irregular white spots with black borders (SMNH-95511#2, Fig. 5 E���F) or a greyish green disc with black rings (SMNH-95511#3, Fig. 5 G���I). Dorsal arms banded light and dark, distal edge of dorsal arm plates with small dark and white spots. The ventral disc is always much lighter than the dorsal disc, often cream with a hint of darker colour, sometimes green. The oral shields are completely creamy white in some specimens. Ventral arms are cream. Some specimens with only up to eight oral papillae. Skeletal elements (Fig. 6) From specimen with 11.7 mm dd (SMHN-178429). Radial shield at widest part about as wide as long, isosceles triangular with abradial projection and convex radial edge, externally with domed center with finer mesh stereom (the domed part is not covered by scales but with granules in the intact animal, all other areas are covered by scales), a circle of pores around domed part, two large pores on the proximal end (Fig. 6A, C). Internally, radial shield with large pore in center of distal half, adjacent to rough patch, distally two knobs, radial one larger, separated by a furrow (Fig. 6B). Dental plates as varying numbers of pieces, sockets for teeth and apical tooth papillae not penetrating, bordered by low ridges, tooth papillae sockets small round, tooth sockets wider oval (Fig. 6D). Adradial genital plate blade-like flat with bulbous distal end and large pore, distal pit with knob on one side, three ridges on other side (Fig. 6 E���F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally (Fig. 6G). Oral plates longer than high, middle part markedly lower than ends, abradial muscle fossa rounded triangular with depression in upper half, not reaching upper edge (Fig. 6H). On adradial oral plate proximoventral part four oral papilla sockets and pores, at proximoventral edge four indistinct granule sockets and pores (Fig. 6I). Vertebrae zygospondylous, with large wing-like muscle fossae (Fig. 6 J���K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening (Fig. 6L, M). Two spurs at proximal external edge and counterparts on internal distal edge of lateral arm plates. Internal lateral arm plate with vertical row of three pores just below the plate centre, fourth pore in tentacle opening edge, bent vertical proximal ridge and small ventral knob, protruding beyond plate edge. Spine articulations inset in the distal plate edge with one thickened lobe, separating muscle and nerve opening. Remarks Ophioderma guineense was until now considered conspecific with O. longicauda. It differs from O. longicauda in having mainly single and flat dorsal arm plates and a higher number of arm spines (Table 1). A previously unrecognized differentiating character is the light colouration of the ventral side of O. guineense, in contrast to the dark colour of O. longicauda, and the spots on the distal edge of the dorsal arm plates, which are absent in O. longicauda. Considering the small morphological differences between the genetic clades in the species complex of O. longicauda, we propose to treat O. guineense as a valid species and we consider the recently collected specimens of genetic cluster C1 as conspecific due to their colouration and mostly single dorsal arm plates. The specimens of NHMD-225525 also concur with this assignment. As lectotype, we selected a specimen that concurs well with the original description (Fig. 4 A���C): Greef (1882) described the colouration of his specimens as on the dorsal disc blackish or dark brown, ventrally light green, the arms dorsally black with grey white rings, ventrally grey yellow, which fits with the material from Senegal (Fig. 5). There are irregular dark rings on the pale brown dorsal disc of the smallest paralectotype examined (Fig. 4G) and tiny dark spots on the dark grey-brown dorsal disc of the lectotype (Fig. 4A), both have cream ventral discs and arms (Fig. 4B, H). The largest of the examined paralectotypes has a faded brown dorsal disc and also some dark brown pigmentation remaining on the ventral disc (Fig. 4 E���F); one of the specimens seen only on photographs has a dark dorsal disc and dark pigment on the ventral disc, quite similar to O. africana sp. nov. (see below, Fig. 7). Based on colour pattern, there may be a possibility that the type specimens are conspecific with O. africana sp. nov. However, in addition to differences in colour, multiple dorsal arm plates are more common in O. africana sp. nov. than in O. guineense, which has led us to assign the Senegal specimens to O. guineense. Colour pattern tends to vary in most species of Ophioderma (Hendler et al. 1995) but O. guineense differs notably from all species observed by this study from Mediterranean and East Atlantic localities (over 800 specimens in the collection of SMNH). These may be similar in dorsal colour with uniformly brown or mottled dorsal discs, sometimes with white spots, but they always have a dark ventral disc and lighter coloured, but never white, arms and oral frames (St��hr et al. 2009). Red pigments seem more common in O. longicauda than green, and the ventral arms are often either reddish or green in the Mediterranean species, with the disc being brown or olive. No embryos have been found in specimens of O. guineense and this species is most likely a broadcast spawner., Published as part of St��hr, Sabine, Weber, Alexandra Anh-Thu, Boissin, Emilie & Chenuil, Anne, 2020, Resolving the Ophioderma longicauda (Echinodermata: Ophiuroidea) cryptic species complex: five sisters, three of them new, pp. 1-37 in European Journal of Taxonomy 600 on pages 13-18, DOI: 10.5852/ejt.2020.600, http://zenodo.org/record/3637773, {"references":["Greeff R. 1882. Echinodermen beobachtet auf einer Reise nach der Guinea-Insel Sao-Thome. Zoologischer Anzeiger 107: 156 - 159.","Koehler R. 1914. Echinoderma I. Asteroidea, Ophiuroidea et Echinoidea. In: Michaelsen E. (ed.) Beitrage zur Kenntnis der Meeresfauna Westafrikas Vol. 1: 127 - 303. L. Friederichsen & Co., Hamburg. https: // doi. org / 10.5962 / bhl. title. 7400","Madsen F. J. 1970. West African ophiuroids. Atlantidae Report 11: 151 - 243.","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 and London.","Stohr S., Boissin E. & Chenuil A. 2009. Potential cryptic speciation in Mediterranean populations of Ophioderma (Echinodermata: Ophiuroidea). Zootaxa 2071: 1 - 20. https: // doi. org / 10.11646 / zootaxa. 2071.1.1"]}

Ophioderma hybrida sp. nov. urn:lsid:zoobank.org:act: 72379251-8181-4B42-8 CB 6-881F0C144526 Figs 9–10; Tables 1–2 Diagnosis Species of Ophioderma with variable morphology. Dorsal arm plates single or as several pieces. Seven to 10 equal arm spines. Eight oral papillae. Colour pattern: disc dark brown or olive dorsally and ventrally, often with white spots. Dorsal arms dark brown with light brown or white and black spots, ventrally lighter brown. Maximum known size about 19 mm dd. Etymology This species has likely evolved by an ancient hybridization event between two species (O. longicauda and O. zibrowii sp. nov. or their respective ancestors), which the Latin word hybrida alludes to. It also refers to the mixed morphology of the species. Material examined Holotype MEDITERRANEAN SEA • Tunisia, Kelibia; 36°50′7.31″ N, 11°7′6.20″ E; depth 0.3–1.5 m; 3 Jul. 2013; Zined Marzouk & Alexandra Weber leg.; preserved in 95% ethanol; SMNH-Type-9185. Paratypes MEDITERRANEAN SEA • 11 specimens; same data as for holotype; SMNH-Type-9186 • 8 specimens; Tunisia, Monastir; 35°46′28.83″ N, 10°50′18.93″ E; depth 2–5 m; 25 Jun. 2013; A. Weber leg.; preserved in 95% ethanol; SMNH-Type-9187. Other material MEDITERRANEAN SEA • 3 specimens (with dorsal disc removed, but present); same data as for holotype; SMNH-178454 • 3 specimens; Tunisia, Tunis; 1849; Åberg leg.; SMNH- 35614, 123374, 123375. Description Holotype (Fig. 9) Disc diameter 17.4 mm, two arms cut off near disc but present (Fig. 9 A–B). Radial shields covered by disc granules (Fig. 9A). Dorsal arm plates weakly tumid, as 1–3 pieces on about first 30 joints, when single twice as wide as long, distal edge straight or concave (Fig. 9 C–D). Ten arm spines, all about equal in length, ventralmost slightly wider. Ventral arm plates overlapping, squared, distal edge convex (Fig. 9E). Two oval scale-like tentacle scales, superimposed on the ventralmost spine. Ventral disc covered by granules similar to dorsal disc. Eight oral papillae, short, weakly pointed to round, Lyman’s ossicle large, pointing into mouth slit (Fig. 9F). Tooth papillae larger than oral papillae, teeth rounded block-shaped. Oral shield almost twice as wide as long, with wide proximal angle, distal edge slightly convex. Madreporite larger, almost as long as wide (Fig. 9B). Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm (Fig. 9F). Adoral shields covered with granules, similar to disc granules. Proximal genital slits angled from oral shield to second arm joint, distal genital slits at disc edge, two arm joints long. Disk dark brown dorsally with radiating lighter brown pattern and scattered white spots (Fig. 9A). Dorsal arms dark brown with light brown and dark spots along the distal plate edges and on some lateral edges (Fig. 9 C–D). Ventral disc and arms light greenish brown, oral shields darker (Fig. 9B). Variation The range of morphological variation is presented in Table 2 and on Fig. 10. The specimens vary in size from 9.8 to 18.7 mm disc diameter. Among the 23 examined specimens, the number of dorsal arm plate pieces varies considerably. Multiple plates or pieces have been found on the proximal arms, until about joint number 30, but not on all joints, and sometimes not on all arms (Fig. 10E). In most specimens, there are only two pieces, but in a few animals, multiple pieces were observed (Fig. 10E, H, N). Some specimens have only single dorsal arm plates (Fig. 10B, K). More pieces are present in the larger specimens but there appears to be only a weak size relation for this trait. Likewise, the number of arm spines and oral papillae is not related to disc diameter. All but two animals have granule covered radial shields. Colour variations are limited. All examined specimens have a dark brown or olive dorsal disc with varying white patterning (from spots to radiating lines) and a dark ventral disc, similar or lighter oral frame and ventral arms. The dorsal arms show alternating light and dark bands and a series of light and dark spots along the distal edge of the dorsal arm plates. Other specimens SMNH- 35614 15.5 mm dd, SMNH-123374 16.3 mm dd, SMNH-123375 14.3 mm dd. Colour faded to light brown. Most of the granules rubbed off the discs, and status of the radial shields as naked/granule covered cannot be assessed, but they have an exposed area, devoid of scales. Eight oral papillae. Two to three dorsal arm plates on single joints (SMNH-123374), up to nine joints (SMNH- 123375), up to 12 joints (SMNH-35614), variable between arms. Juveniles present in the bursae. Remarks The origin of this species through an ancient hybridization event between O. longicauda and O. zibrowii sp. nov. (or their ancestors) (Weber et al. 2019) is further supported by its high morphological variability. Multiple dorsal arm plates are typical of O. longicauda, and in some specimens of O. hybrida sp. nov. the most proximal joints bear several plates, forming a crowded mosaic pattern similar to O. longicauda. The usually granule covered radial shields are shared with all species except O. zibrowii sp. nov. (Table 1) and are probably evidence of O. longicauda (or its ancestor) being one of the ancestral species. The variation of arm spines and oral papillae overlap with both ancestral species and O. africana sp. nov., but O. guineense has a higher number of arm spines. Ophioderma hybrida sp. nov. shares the vivid colour pattern of the arms with O. zibrowii sp. nov. and O. guineense, whereas O. africana sp. nov. and O. longicauda are darker and the spots on the arms are less obvious in the former. So far, O. hybrida sp. nov. appears to be a smaller species with disc diameter under 20 mm, but more material is needed to confirm this. We believe that the reproductive mode of O. hybrida sp. nov. is brooding given that brooding individuals have been collected in Tunisia (Stöhr et al. 2009) and the ones examined in this study are morphologically more similar to O. hybrida sp. nov. than to O. zibrowii sp. nov., because they have more dorsal arm plates per joint. Yet, we cannot confirm it, as the brooding specimens from Tunisia were too old for genetic analyses (1849 and 1924; PCR amplification failed). In addition, all other O. hybrida sp. nov. specimens have been collected outside the reproductive season so examination of their gonads was inconclusive in determining their reproductive strategy.

Ophioderma zibrowii sp. nov. urn:lsid:zoobank.org:act: 7FB245FD-C8DF-402A-A7B1-FA7D048AFB04 Figs 11���12; Table 1 Diagnosis Usually naked radial shields. Seven arm spines. Eight oral papillae. Colour pattern greenish brown to dark green variegated on dorsal and ventral disc, arms banded, with light and dark spots along distal dorsal plate edges, oral frame and ventral arms lighter. Largest known size 21 mm dd. Brooder. Etymology The species is named after Helmut Zibrowius for his important contributions in collecting material for this study. The existence of brooding Ophioderma in the Mediterranean Sea might still be unknown without his effort. Material examined Holotype MEDITERRANEAN SEA ��� ♀ (brooding); southern Crete, Agios Pavlos; May 2012; A. Weber leg.; preserved in 95% ethanol; SMNH-Type-9188. Paratypes MEDITERRANEAN SEA ��� 1 ♀ (brooding); same data as for holotype; SMNH-Type-9189 ��� 7 specimens; Greece, Symi Island, near Rhodes, Agios Georgios; 10 Aug. 2005; R. Graille leg.; preserved in 95% ethanol; SMNH-Type-9190 to 9196. Other material MEDITERRANEAN SEA ��� 18 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, northwestern Crete, Elounda, Plaka Beach; 2 Jul. 2008; Emilie Aegea leg.; preserved in 95% ethanol; SMNH-102712, 113690 to 13697, 11370 to 113704, 113709 to 113712 ��� 21 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, Rhodes, Kallithea Bay; 9 Sep. 2005; H. Zibrowius leg.; preserved in 95% ethanol; SMNH-99657, 113982, 113985, 113987 to 113990, 113992 to 114003, 114005, 114006 ��� 41 specimens (mitochondrial COI lineage L3, nuclear cluster C5); Greece, Crete; May 2012; A. Weber leg.; preserved in 95% ethanol; SMNH-147070 ��� 1 specimen (dd = 15.4 mm), dried dissociated ossicles and ossicles on two SEM stubs; same data as for preceding; SMNH-178455. Description Holotype (Fig. 11) Disc diameter 14.3 mm, one intact arm, all others removed for molecular analysis. Dorsal disc with dense cover of granules, radial shields naked, widely separated, visible part oval (Fig. 11A). Granules and scales crowd onto basal arms. Dorsal arm plates single on all joints, flat, overlapping, broadly fanshaped, twice as wide as long, with straight edges, wider distally than proximally (Fig. 1D). Up to seven conical, slightly pointed arm spines, about equal in length, half as long as arm joint, ventralmost spine thicker than others and blunt (Fig. 7C). Ventral disc covered by granules similar to dorsal disc. Adoral shields and oral plates covered by granules, oral shields naked (Fig. 11B). Seven to eight oral papillae, Lyman���s ossicle elongated, pointing into mouth slit, adoral shield spines two, block shaped, other papillae smaller, conical (Fig. 11E). Tooth papillae larger, conical, two in line with oral papillae. Oral shields triangular, twice as wide as long, straight distal edge, obtuse rounded proximal angle. Madreporite as wide as long. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules, similar to disc granules. Proximal genital slits parallel to and as long as first and second joints. Distal genital slit parallel to and as long as fourth joint, at distance from disc edge. Bursae filled with brooded juveniles (Fig. 11F). Dorsal disc slightly variegated dark brown-green with grey, dorsal arms greyish green with brown bands, each light and dark band about three joints wide, a series of larger white and smaller black spots along the distal edge of each dorsal arm plate (Fig. 11A, D). Ventral disc as dark as dorsal, oral frame and ventral arms lighter grey-green (Fig. 11B). Arm spines grey-green with white tips, except ventralmost spine, which is white (Fig. 11 C���D). Oral papillae and tentacle scales white (Fig. 11C, E). Paratypes Disc diameter 9.7���13.5 mm.All have up to seven arm spines. Most paratypes have a green colour pattern similar to the holotype, one specimen is olive-brown. One paratype with granule-covered radial shields, all others naked. Oral papillae eight, in two specimens nine, but variable between jaws in an individual. Two specimens with two dorsal arm plates on 2���3 proximal joints on 1���3 arms. One specimen with 2���3 dorsal arm plates on proximal joints on three arms, on seven joints on one of those arms. One specimen has a parasitic snail (Ersilia Monterosato, 1872 sp.) on one arm. All paratypes lack part of an arm, which was used for molecular analysis. Non-paratypes (SMNH-147070) Disc diameter 8.4���20.5 mm. Specimens smaller than 10 mm dd had six arm spines, all others seven. Most specimens have a green colour pattern similar to the holotype, several were dark olive with numerous white granules scattered across the disc. Out of the 42 specimens, a single one had granule-covered radial shields, all others with naked radial shields. Most of the specimens above 10 mm dd had at least two proximal arm joints on at least one arm with two dorsal arm plates, up to seven joints and several arms with double plates in one animal. The plates were usually of equal size or there was a hairline crack along the midline of the plate. Rarely, a third dorsal arm plate was present on one or two joints. Oral papillae eight, rarely nine, but variable between jaws in an individual and not correlated to size. These specimens were used for thermotolerance and regeneration experiments. These specimens were kept in an aquarium for a year where experiments of thermotolerance and regeneration were conducted. Therefore, most arms show signs of regeneration. One specimen had bright white distal arms but no sign of healed breaks. Maximum size recorded for O. zibrowii sp. nov. was 21 mm dd (Weber et al. 2014). Skeletal elements (Fig. 12) Non-type specimen of 15.4 mm dd (SMNH-178455), completely dissociated. Radial shields elongated isosceles triangular, external surface with embossed oval of more finely meshed stereom in center of domed and thickened proximal half (only visible part in intact skeleton), row of eight pores curves around proximal dome (Fig. 12A, C). On inner side of radial shield, a large pore on distal half and a round patch with denser stereom, two low distal bulbs separated by a furrow (Fig. 12B). Dental plate consists of several pieces, sockets not penetrating the plate, bordered by low rim, tooth papillae sockets smaller, round, tooth sockets wide, oval (Fig. 12D). Adradial genital plate blade-like flat with bulbous distal end and large pore, distal pit with knob on one side, three ridges on other side (Fig. 12 E���F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally (Fig. 12G). Oral plates longer than high, middle part markedly lower than ends, abradial muscle fossa rounded triangular, not reaching upper edge (Fig. 12H). On adradial oral plate proximoventral part four oral papilla sockets and pores, at proximoventral edge four granule sockets and pores (Fig. 12I). Vertebrae typically zygospondylus, with large dorsal and small ventral muscle flanges, no growth rings obvious (Fig. 12 J���K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening (Fig. 12 L���M). Two spurs at proximal external edge and counterparts on internal distal edge of lateral arm plates. Internal lateral arm plate with vertical row of three pores just below the plate centre, distal to curved vertical ridge, fourth pore in tentacle opening edge, small ventral knob, protruding beyond plate edge. Spine articulations inset in the distal plate edge with one thickened lobe, separating muscle and nerve opening. Remarks The species determination rests mainly on the genetic analyses. We chose the predominant genetic group (C5) to describe a new species, but due to its short reproduction period (May���June, Weber et al. 2014) most specimens of this cluster happen to have been collected outside the brooding time. Since brooding is one of the key characters of this species, we felt that the holotype should be a brooding female. It is not ideal that the chosen animal has only a single intact arm remaining, but in a pentaradially symmetric animal one arm is sufficient and representative of the others. Morphological differentiation from O. longicauda rests on the usually naked radial shields and the single, flat arm plates. Also, O. zibrowii sp. nov. usually lacks red pigments which are common in O. longicauda, whereas green pigments are common in O. zibrowii sp. nov. but absent in O. longicauda. In addition, O. zibrowii sp. nov. commonly has a series of white and black spots along the distal edge of the dorsal arm plates, which are shared with O. hybrida sp. nov. and O. guineense. The radial shields differ microscopically from those of O. longicauda in the clear demarcation of an embossed part with finer meshed stereom, representing the naked visible part of the radial shields in the intact animal. The major part of the radial shield is overlapped and covered by disc scales, obscuring its true shape. Possibly, the more elongated oral plates and higher lateral arm plates in O. zibrowii sp. nov. distinguish the species, but these small differences need to be studied with morphometric methods and larger samples. Brooding females have been collected from Cyprus and Lebanon, but they belong to another genetic cluster (C6). Morphologically they cannot be distinguished at this time and we consider them to be allopatric populations within the same species. They are excluded from the concept of the new species O. zibrowii sp. nov., as new evidence may in the future lead to them being recognized as a different species., Published as part of St��hr, Sabine, Weber, Alexandra Anh-Thu, Boissin, Emilie & Chenuil, Anne, 2020, Resolving the Ophioderma longicauda (Echinodermata: Ophiuroidea) cryptic species complex: five sisters, three of them new, pp. 1-37 in European Journal of Taxonomy 600 on pages 28-32, DOI: 10.5852/ejt.2020.600, http://zenodo.org/record/3637773, {"references":["Weber A. A. - T., Stohr S. & Chenuil A. 2014. Genetic data, reproduction season and reproductive strategy data support the existence of biological species in Ophioderma longicauda. Comptes Rendus Biologies 337: 553 - 560. https: // doi. org / 10.1016 / j. crvi. 2014.07.007"]}

Ophioderma africana sp. nov. urn:lsid:zoobank.org:act: 64021157-EA56-449 B- 9BCB-FD02F10CC67F Figs 7���8; Table 1 Diagnosis Mostly single dorsal arm plates, but several plates on some joints, eight arm spines, eight oral papillae. Colour dark brown dorsally, ventral disc as dark, ventral arms lighter brown, black and white spots along distal edge of dorsal arm plates. Maximum size under 19 mm dd (26 mm dd if non-type material is considered). Etymology The species is named for its area of collection, off (West) Africa. Material examined Holotype TROPICAL ATLANTIC OCEAN ��� Senegal, Dakar, Gor��e Island, shipwreck Tacoma; depth 10 m; 28 Aug. 2007; H. Zibrowius leg.; scuba diving, hand collected; under stones; preserved in 95% ethanol; SMNH-Type-7484. Paratypes TROPICAL ATLANTIC OCEAN ��� 22 specimens (6 sampled for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); same data as for holotype; SMNH-Type-7485 ��� 1 specimen, ossicles, 4 remaining arms, 2 SEM stubs with ossicles; same data as for holotype; dried; SMNH-Type-7486 ��� 11 specimens (including a separate vial with eggs that were spawned during collection); same data as for holotype; initially in formalin, then 80% ethanol; SMNH-Type-7487 ��� 4 specimens (2 used for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); Senegal, Dakar, Cap Manuel; depth 7 m; 6 Sep. 2007; H. Zibrowius leg.; under stones; scuba diving, hand collected; preserved in 95% ethanol; SMNH-Type-7488 ��� 4 specimens (2 used for both COI and nuclear markers, mitochondrial COI lineage L 5, nuclear cluster C 2); Senegal, Madelene Islands; depth 10���12 m; 9 Sep. 2007; H. Zibrowius leg.; among piled up rocks; scuba diving, hand collecting; preserved in 95% ethanol; SMNH-Type-9184. Other material TROPICAL ATLANTIC OCEAN ��� 6 specimens (labelled as O. longicauda); Senegal, between Gor��e and Tiaroye; depth 15���20 m; Feb.���Mar. 1952; Thorson leg.; preserved in ethanol; NHMD-225514 ��� 6 specimens (labelled as O. longicauda); Senegal, Dakar; Jan. 1928; Sudan exped.; H. Madsen leg.; dried; partial sample; NHMD- 225517 ��� 10 specimens (labelled as O. longicauda); Dakar; 6 Jan. 1928; Sudan exped.; H. Madsen & O. Hagerup leg.; preserved in in ethanol; partial sample; NHMD-225523 ��� 1 specimen (labelled as O. longicauda); Senegal, Dakar, rocky coast; 3 Jan. 1928; H. Madsen leg.; preserved in ethanol; NHMD-225524. Description Holotype (Fig. 7) Disc diameter 18.7 mm, disc completely covered by small granules dorsally and ventrally (Fig. 7 A���C), including the radial shields (Fig. 7B). Disc scales with granules crowding onto arm base, covering first dorsal arm plate except for a mid-distal portion, second plate also covered by scales and granules laterally (Fig. 7 D���E). Dorsal arm plates flat, 2.5 times as wide as long, with straight distal edge, contiguous, mostly single, but joints with two plates common (often unequal in size) (Fig. 7 D���E). Eight arm spines, ventralmost one thicker and slightly longer than others (Fig. 7F). Ventral disc covered by granules similar to dorsal disc. Eight conical oral papillae, Lyman���s ossicle and adoral shield spines hardly distinguishable (Fig. 7G). Two larger tooth papillae at jaw tips in line with oral papillae, blocklike teeth. Oral shields rounded triangular, slightly wider than long, madreporite slightly wider than other oral shields and with central round depression. Adoral shield long, narrow, extending around the lateral edge of the oral shield, separating it from the arm. Adoral shields covered with granules, similar to disc in size. Proximal genital slits about one joint long, distal slits about three joints long (Fig. 7C). Ventral arm plates contiguous, as wide as long, with wide proximal angle, convex distal edge (Fig. 7F). Two oval scale-like tentacle scales, superimposed on the ventralmost spine. Disc dark brown dorsally, with some lighter brown patches. Dorsal arms dark brown, distally lighter and banded. Dorsal arm plates with small light brown and black dots along distal edge. Ventral disc as dark as dorsal, oral frame and ventral arms light orange brown (Fig. 7A, C). Paratypes Size range 12���18.4 mm dd. The specimens have dark brown to almost black dorsal discs (on some individuals with lighter patches), arms with dark and lighter bands. Ventral discs as dark as dorsal, oral frames and ventral arms lighter brown (Fig. 7H). Individuals with small light spots on the disc occur. Most specimens with black dots on the distal edge of the dorsal plates, but some without. Dorsal arm plates are mostly single, but most arms with several joints with two plates. Skeletal elements (Fig. 8) From specimen of 15.7 mm dd (SMNH-Type-7486) with mostly single dorsal arm plates but some small additional plates on few joints. Seven oral papillae, all similar except Lyman���s ossicle. Up to eight arm spines. Radial shields isosceles triangular with deeply concave abradial edge and abradial distal processes, distal center domed with small pores (the domed part is not covered by scales but with granules in the intact animal, all other areas are covered by scales) (Fig. 8A, C). Inner side of radial shield, on the distal articulation surfaces, two larger oval depressions and a smaller round one between these (Fig. 8B). Dental plate consists of several pieces with non-perforating sockets for 2���3 tooth papillae and single wide teeth, granule pores not obvious (Fig. 8D). Adradial genital plate blade-like flat with bulbous distal end and large pore, distal pit with knob on one side, two distal depressions on other side (Fig. 8 E���F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally (Fig. 8G). Oral plates longer than high, middle part slightly lower than ends, adradially with four oral papilla pores on ventral edge of proximal part, small granule pores at ventral edge (Fig. 8I). Adradial distal muscle fossa wide ventrally, tapering dorsalwards, shorter than plate height, articulation area slightly higher than wide (Fig. 8H). Suture line diagonally across oral plate mid-line, separated easily during preparation on several jaws. Vertebrae typically zygospondylus, with large dorsal and small ventral muscle flanges, no growth rings obvious (Fig. 8 J���K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening (Fig. 8 L���M). Two spurs at proximal external edge and counterparts on internal distal edge of lateral arm plates. Internal lateral arm plate with vertical row of three pores just below the plate centre, fourth pore in tentacle opening edge, bent vertical proximal ridge and small ventral knob, protruding beyond plate edge. Spine articulations inset in the distal plate edge with one thickened lobe, separating muscle and nerve opening. Other specimens The specimens from NHMD possess mostly single dorsal arm plates, on few joints there are more than one and some specimens have on the most proximal joints a mosaic of slightly tumid plates. Some specimens have naked radial shields. They have 7���9 oral papillae, among which the adoral shield spine and Lyman���s ossicle are sometimes obvious, differing in shape from the other papillae. They are uniformly dark brown or variegated dark, on both dorsal and ventral sides. They range in size between 19 and 26 mm dd and are assigned to O. africana sp. nov. mostly by colour pattern and exclusion of O. longicauda from off West Africa. The specimen NHMD-225524, which measures only 5.7 mm dd, has a light brown dorsal pattern with faint spots, a cream white ventral surface, and single dorsal arm plates, similar to O. guineense and is reassigned to that species. Remarks Ophioderma africana sp. nov. shares the dark colour on both sides of the disc with O. longicauda, its spine numbers fall within the variability of both O. longicauda and O. guineense, it has as few oral papillae as O. longicauda (Table 1). These characters are however variable and small specimens may be difficult to identify. Nuclear and mitochondrial data were distinct from O. longicauda and all other clusters found (Weber et al. 2019), placing O. africana sp. nov. in cluster C2 and COI lineage L5. Some of the animals started spawning during collecting, which indicates that the species is a broadcast spawner, not a brooder., Published as part of St��hr, Sabine, Weber, Alexandra Anh-Thu, Boissin, Emilie & Chenuil, Anne, 2020, Resolving the Ophioderma longicauda (Echinodermata: Ophiuroidea) cryptic species complex: five sisters, three of them new, pp. 1-37 in European Journal of Taxonomy 600 on pages 19-22, DOI: 10.5852/ejt.2020.600, http://zenodo.org/record/3637773, {"references":["Weber A. A. - T., Stohr S. & Chenuil A. 2019. Species delimitation in the presence of strong incomplete lineage sorting and hybridization: lessons from Ophioderma (Ophiuroidea: Echinodermata). Molecular Phylogenetics and Evolution 131: 138 - 148. https: // doi. org / 10.1016 / j. ympev. 2018.11.014"]}