Stöhr, 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, European Journal of Taxonomy 600, pp. 1-37: 28-32
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Ophioderma zibrowii sp. nov.
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.
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.
MEDITERRANEAN SEA • ♀ (brooding); southern Crete, Agios Pavlos ; May 2012; A. Weber leg.; preserved in 95% ethanol; SMNH-Type-9188.
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 .
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 .
Holotype ( Fig. 11View Fig)
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. 11AView Fig). 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. 1DView Fig). 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. 7CView Fig). Ventral disc covered by granules similar to dorsal disc. Adoral shields and oral plates covered by granules, oral shields naked ( Fig. 11BView Fig). Seven to eight oral papillae, Lyman’s ossicle elongated, pointing into mouth slit, adoral shield spines two, block shaped, other papillae smaller, conical ( Fig. 11EView Fig). 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. 11FView Fig).
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, DView Fig). Ventral disc as dark as dorsal, oral frame and ventral arms lighter grey-green ( Fig. 11BView Fig). Arm spines grey-green with white tips, except ventralmost spine, which is white ( Fig. 11View Fig C–D). Oral papillae and tentacle scales white ( Fig. 11C, EView Fig).
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.
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. 12View Fig)
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, CView Fig). 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. 12BView Fig). 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. 12DView Fig). 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. 12View Fig E–F). Abradial genital plate much smaller and scale-like flat, distally wider than proximally ( Fig. 12GView Fig). Oral plates longer than high, middle part markedly lower than ends, abradial muscle fossa rounded triangular, not reaching upper edge ( Fig. 12HView Fig). On adradial oral plate proximoventral part four oral papilla sockets and pores, at proximoventral edge four granule sockets and pores ( Fig. 12IView Fig). Vertebrae typically zygospondylus, with large dorsal and small ventral muscle flanges, no growth rings obvious ( Fig. 12View Fig J–K). Lateral arm plates compact, weakly curved around the arm, strongly convex distally, concave proximally, with ventral excavation for tube foot opening ( Fig. 12View Fig 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.
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.
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