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Genus Ospreyella L��ter & W��rheide (in L��ter et al. 2003) Type species. Ospreyella depressa L��ter (in L��ter et al. 2003), 2003 by original designation. Emended diagnosis. See Simon & Hoffmann (2013, p. 412)., Published as part of Simon, Eric, Hiller, Norton, Logan, Alan, Theuerkauff, Dimitri & Mottequin, Bernard, 2019, Recent thecideide brachiopods from a submarine cave in the Department of Mayotte (France), northern Mozambique Channel, pp. 201-239 in Zootaxa 4613 (2) on page 214, DOI: 10.11646/zootaxa.4613.2.1, http://zenodo.org/record/3238856, {"references":["Luter, C., Worheide, G. & Reitner, J. (2003) A new thecideid genus and species (Brachiopoda, Recent) from submarine caves of Osprey Reef (Queensland Plateau, Coral Sea, Australia). Journal of Natural History, 37, 1423 - 1432. http: // doi. org / 10.1080 / 00222930110120971","Simon, E. & Hoffmann, J. (2013) Discovery of Recent thecideide brachiopods (Order: Thecideida, Family: Thecideidae) in Sulawesi, Indonesian Archipelago, with implications for reproduction and shell size in the genus Ospreyella. Zootaxa, 3694 (5), 401 - 433. http: // doi. org / 10.11646 / zootaxa. 3694.5.1"]}

Genus Thecidellina Thomson, 1915 Type species. Thecidium barretti Davidson, 1864 by original designation., Published as part of Simon, Eric, Hiller, Norton, Logan, Alan, Theuerkauff, Dimitri & Mottequin, Bernard, 2019, Recent thecideide brachiopods from a submarine cave in the Department of Mayotte (France), northern Mozambique Channel, pp. 201-239 in Zootaxa 4613 (2) on page 205, DOI: 10.11646/zootaxa.4613.2.1, http://zenodo.org/record/3238856, {"references":["Thomson, J. A. (1915) A new genus and species of Thecidiinae. Geological Magazine, New Series, Decade VI, 2, 461 - 464. https: // doi. org / 10.1017 / S 0016756800203531","Davidson, T. (1864) On the recent and Tertiary species of the genus Thecidium. Geological Magazine, 1, 12 - 22. http: // doi. org / 10.1017 / s 0016756800469803"]}

Genus Minutella Hoffmann & L��ter, 2010 Type species. Minutella tristani Hoffmann & L��ter, 2010, by original designation., Published as part of Simon, Eric, Hiller, Norton, Logan, Alan, Theuerkauff, Dimitri & Mottequin, Bernard, 2019, Recent thecideide brachiopods from a submarine cave in the Department of Mayotte (France), northern Mozambique Channel, pp. 201-239 in Zootaxa 4613 (2) on page 211, DOI: 10.11646/zootaxa.4613.2.1, http://zenodo.org/record/3238856, {"references":["Hoffmann, J. & Luter, C. (2010) Shell development in thecidellinine brachiopods with description of a new Recent genus. Special Papers in Palaeontology, 84, 137 - 160."]}

Comparison with Thecidellina europa Logan et al., 2015 In this species, the ventral valve floor has a wider median area between gonad pits that is ornamented with smaller tubercles. The dorsal median septum is thicker, the marsupial orifices are smaller and differently shaped. Canopying spicules are thicker at their base near the valve floor and fuse together producing finely granulate canopies with irregular ventral surfaces covering two-thirds of the brachial cavities (Logan et al. 2015, fig. 1L)., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 494, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Logan, A., Hoffmann, J. & Luter, C. (2015) Checklist of Recent thecideoid brachiopods from the Indian Ocean and Red Sea, with a description of a new species of Thecidellina from Europa Island and a re-description of T. blochmanni Dall from Christmas Island. Zootaxa, 4013 (2), 225 - 234. https: // doi. org / 10.11646 / zootaxa. 4013.2.4"]}

Comparison with Thecidellina congregata Cooper, 1954 1954 Thecidellina congregata: Cooper, pp. 316���317; pl. 80, figs. 8���13. 1983 Thecidellina congregata: Grant, pp. 82���83; figs. 6���8. 2008 Thecidellina congregata: Logan, pp. 411���412, figs. 6.10���6.17. 2007 Thecidellina maxilla: Bitner, fig. 3F. 2015 Thecidellina maxilla: Bitner, fig. 5A���E. The adult shell is larger. Brachial cavities have peculiar single canopying spicules that are very massive in the anterior part of the canopies and much thinner in their posterior part (Cooper 1954, pl. 80, figs. 10���13; Logan, 2008, figs. 6���17). In the smaller shell of T. mawaliana sp. nov. the canopies are made of very thin, fragile spicules all fused together., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 494, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Cooper, G. A. (1954) Recent brachiopods in Bikini and nearby atolls, Marshall Islands. United States Geological Survey Professional Paper, 260 G, 315 - 318. https: // doi. org / 10.3133 / pp 260 g","Logan, A. (2008) Holocene thecideide brachiopods from the north-western Pacific Ocean: systematics, life habits and ontogeny. Systematics and Biodiversity, 6 (3), 405 - 413. https: // doi. org / 10.1017 / s 1477200008002739"]}

Comparison with Thecidellina maxilla (Hedley, 1899) 1899 Thecidea maxilla: Hedley, pp. 508���510, fig. 57. 1920 Thecidellina maxilla: Dall, p. 283. 1927 Thecidellina maxilla: Thomson, p. 140. 1954 Thecidellina maxilla: Cooper, pp. 317���318, pl. 81, figs. 1���10. 1985 Thecidellina maxilla: Zezina, p. 208. non 1987 Thecidellina maxilla: D���Hondt, p. 41, pl. 4, figs. 1���5. ? 1997 Thecidellina maxilla: Laurin, pp. 453���454, fig. 40A���B. pp 2007 Thecidellina maxilla: Bitner, p. 498, fig. 3A���E, 3G���H, non fig. 3F. non 2009 Thecidellina maxilla: Bitner, pp. 17���18, fig. 12A���J. pp 2014 Thecidellina maxilla: Bitner, p. 259, fig. 11F���I (non fig. 11D���E). non 2015 Thecidellina maxilla: Bitner, p. 42, fig. 5A���E. T. maxilla is a large species with horse-shoe shaped dorsal valve with a peripheral ridge smooth to finely granulate. T. mawaliana sp. nov. is a rather small species drop-like shaped with a peripheral ridge ornamented with strong tubercles. In T. maxilla the brachial bridge is relatively narrower (Hedley 1899, fig. 57) and the median septum is long, with lateral flanges developed along the inner sides of the brachial cavities (Hedley 1899, fig. 57, p. 509; Cooper 1954, pl. 81, fig. 10). The brachial cavities are more elongate. The single massive spicules of the intrabrachial ridge which are not completely covering the brachial cavities are rather regular in shape and size. In T. mawaliana sp. nov. the brachial bridge is wider and the median septum is devoid of lateral flanges. The brachial cavities are oval. The spicules of the brachial cavities, forming complete canopies are thin, irregular in shape and fused together. The ventral valve floor in T. maxilla is not roughly tuberculate (nearly smooth), the planodeltidium is larger and quite concave dorsally (Cooper 1954, pl. 81, figs. 5���6). The hemispondylium in T. maxilla is not attached to the valve floor. Remarks. Non T. maxilla in D���Hondt (1987). In this paper, this species is cited without description but the illustrated specimens are very distinct from the type of T. maxilla. Their median septum is extremely wide and the spicules, differently shaped, form complete canopies on the brachial cavities. This is possibly a new species and this material must be restudied to give accurate taxonomical data. ? T. maxilla in Laurin (1997). This identification remains unclear as no detailed description is provided and the illustrations in this paper are not informative. This material must be revisited to have accurate data for determination. Non T. maxilla in Bitner (2007, p. 498, fig. 3F). This specimen is probably a representative of T. congregata Cooper, 1954. Canopying spicules are different from the type of T. maxilla, being single and very massive in the anterior part of the brachial cavities and being extremely small and pointed in the posterior part. In T. maxilla, spicules have a much more regular shape. The right and left sides of the septum have no typical ���curled flanges with serrate edges��� as in T. maxilla (Hedley 1899, fig. 57; Cooper 1954, pl. 81, fig. 10). Non T. maxilla in Bitner (2009, pp. 17���18; fig. 12A���J). These specimens display a peripheral ridge with strong tubercles and a median septum without lateral flanges. In T. maxilla the peripheral ridge is nearly smooth or finely granulated. The space between the brachial bridge and intrabrachial ridge is larger than in T. maxilla. The ventral valve is not emarginate as in the type specimen and has a rough valve floor whereas it is nearly smooth in T. maxilla. The cardinal process and socket ridges are much stronger than in the type population of T. maxilla. These specimens could be representatives of another species derived from the same group as T. maxilla but it needs genetic confirmation. Non T. maxilla in Bitner (2014, p. 259, fig. 11D���E). These ventral valves have a floor with extremely thick tubercles whereas this surface in the T. maxilla is smooth or finely granulate. The gonad pits are situated more posteriorly and they are much smaller than in T. maxilla or in T. mawaliana sp. nov. Moreover, it has a narrow median groove devoid of tubercles (not a ridge), a character not seen in T. maxilla. They are also not similar to T. mawaliana sp. nov.: in the new species, the valve floor displays much smaller tubercles, the hemispondylium is attached to the bottom of the valve floor for only a short part of its length and the gonad pits are larger. Non T. maxilla in Bitner (2015, fig. 5A���E). Canopying spicules are different from the type of T. maxilla, being single and very massive in the anterior part of the brachial cavities and being extremely small and pointed in the posterior part. The typical ���curled flanges with serrate edges��� of the median septum are missing. These specimens are representatives of T. congregata (see hereunder)., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on pages 492-493, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Hedley, C. (1899) The Mollusca of Funafuti. Part II. Pelecypoda and Brachiopoda. Memoirs of the Australian Museum, 3, 491 - 510. https: // doi. org / 10.3853 / j. 0067 - 1967.3.1899.504","Cooper, G. A. (1954) Recent brachiopods in Bikini and nearby atolls, Marshall Islands. United States Geological Survey Professional Paper, 260 G, 315 - 318. https: // doi. org / 10.3133 / pp 260 g","Laurin, B. (1997) Brachiopodes recoltes dans les eaux de la Nouvelle-Caledonie et des iles Loyaute, Matthew et Chesterfield. In: Crosnier, A. (Ed.), Resultats des Campagnes MUSORSTOM, volume 18. Memoires du Museum national d'Histoire Naturelle, 176, 411 - 471.","Bitner, M. A. (2007) Recent brachiopods from the Austral Islands, French Polynesia, South-Central Pacific. Zoosystema, 29 (3), 419 - 461.","Bitner, M. A. (2009) Recent Brachiopoda from the Norfolk Ridge, New Caledonia, with description of four new species. Zootaxa, 2235, 1 - 39.","Bitner, M. A. (2014) Living brachiopods from French Polynesia, Central Pacific, with descriptions of two new species. Pacific Science, 68 (2), 245 - 265. https: // doi. org / 10.2984 / 68.2.6","Bitner, M. A. (2015) Checklist of recent brachiopod species collected during the Terrasses and Exbodi cruises in the New Caledonian region, SW Pacific. ZooKeys, 537, 33 - 50. https: // doi. org / 10.3897 / zookeys. 537.6567"]}

Comparison with Thecidellina insolita Hoffmann et al., 2009 In T. insolita the cardinal process has a wider median spur than in T. mawaliana sp. nov. The calcitic pole is not connected to the valve floor in T. insolita whereas it is strongly fused with the valve floor in T. mawaliana sp. nov. In T. insolita the brachial cavities have single massive canopying spicules and marsupial notches are larger and more quadrangular., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 494, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832

Ospreyella mutiara Simon & Hoffmann, 2013 Text-Fig. 2; Pl. 7, Figs. 3a���d, 4a���f 2013 Ospreyella mutiara: Simon & Hoffmann, pp. 412���424, text-fig. 4, pl. 3, figs. 1���8, pl. 4., figs. 1���10, pl. 5, figs. 1���5, pl. 6, figs. 1���8, pl. 7, figs. 1���6. 2016 Ospreyella mutiara: Simon et al., p. 5. Material. Available specimens were collected in the shipwreck from sieved sediment or attached on oyster shells and preserved in ethanol for a future DNA study. This species is quite scarce at this station and the state of preservation is not excellent. The dried material is represented by ten individuals: eight articulated shells, one dorsal valve and two ventral valves. Material preserved in ethanol represents six specimens supposed to be alive when collected and are reserved for future DNA analyses. Description. The material collected in Lembeh corresponds completely to the diagnosis and detailed description given for this species in 2013 by Simon & Hoffmann (pp. 412���415). The material from the Strait of Lembeh is thus similar to the material that was found in Donggala in the Strait of Makassar. The only other known possible Ospreyella species could be O. palauensis Logan, 2008. However, the specific characters of the latter species are not observed in O. mutiara from Lembeh, which has a median depression strongly tuberculate in its anterior part, a narrow ventral surface of the thick median ramus filled with secondary material, thick ramuli as wide as ramus filled with secondary material and frilled, minor interbrachial lobes asymmetrical and subparallel. A complete comparison between O. mutiara and O. palauensis can be found in Simon and Hoffmann (2013, tab. 4, p. 423). Ontogeny and molecular analyses. The developments of the shell and of the lophophore have been described, discussed and illustrated in detail by Simon & Hoffmann (2013, pp. 427���428, pl. 6, figs. 1���8, pl. 7, figs. 1���6). In the same paper (p. 428, text-fig. 4) results of a molecular analyze of 18S rDNA has been published for O. mutiara that confirmed the Indo-Pacific distribution for the genus and the validity of the specific status of this species., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 496, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Simon E. & Hoffmann, J. (2013) Discovery of Recent thecideide brachiopods (Order: Thecideida, Family: Thecideidae) in Sulawesi, Indonesian Archipelago, with implications for reproduction and shell size in the genus Ospreyella. Zootaxa, 3694 (5), 401 - 433. https: // doi. org / 10.11646 / zootaxa. 3694.5.1","Logan, A. (2008) Holocene thecideide brachiopods from the north-western Pacific Ocean: systematics, life habits and ontogeny. Systematics and Biodiversity, 6 (3), 405 - 413. https: // doi. org / 10.1017 / s 1477200008002739"]}

Thecidellina mawaliana sp. nov. Table 1; Text-Figs. 2���4; Pls. 1���6 Holotype: RBINS���BT.9 (Pl. 1, Fig. 2a���g, Pl. 5, Fig. 4): articulated adult specimen. Paratypes: RBINS���BT.10 (Pl. 1, Fig. 1a���f): articulated adult specimen. RBINS���BT.11 (Pl. 3, Fig. 1a���n): adult dorsal valve with complete development of the canopies on brachial cavities. RBINS���BT.12 (Pl. 2, Fig. 1a���g): small articulated specimen with brachial cavities still without canopying spicules and with a planodeltidium curving to right side. RBINS���BT.13 (Pl. 2, Fig. 2a���g, Pl. 4, Fig. 2a���c): juvenile articulated specimen. RBINS��� BT.14 (Pl. 2, Fig. 3): small articulated specimen with its planodeltidium curving to the left side. RBINS���BT.15 (Pl. 3, Fig. 2a���f): adult ventral valve with teeth and intact hemispondylium. RBINS���BT.16 (Pl. 4, Fig. 1a���d): a very early juvenile dorsal valve. RBINS���BT.17 (Pl. 4, Fig. 3a���f): juvenile dorsal valve with a peribrachial ridge beginning its development and with appearing posterior outgrowths from the brachial bridge. RBINS���BT.18 (Pl. 4, Fig. 4a���c): juvenile dorsal valve with tubercles beginning the building of the brachial cavities. RBINS���BT.19 (Pl. 5, Fig. 1a���g): dorsal valve with developed median septum, first appearance of calcitic pole and continuing the development of brachial cavities. RBINS���BT.20 (Pl. 5, Fig. 2a���e): dorsal valve with brachial bridge and peribrachial ridge connected. RBINS���BT.21 (Pl. 5, Fig. 3): adult dorsal valve with brachial cavities with incomplete canopies. RBINS���BT.22 (Pl. 5, Fig. 6), RBINS���BT.23 (Pl. 5, Fig. 7), RBINS���BT.24 (Pl. 5, Fig. 5a���b), and RBINS���BT.25 (Pl. 5, Fig. 8a���b): several adult dorsal valves illustrating the variation of the canopies covering the brachial cavities. RBINS���BT.26 (Pl. 6, Fig. 1), RBINS���BT.27 (Pl. 6, Fig. 2), RBINS���BT.28 (Pl. 6, Fig. 3), RBINS���BT.29 (Pl. 6, Fig. 4), RBINS���BT.30 (Pl. 6, Fig. 5), and RBINS���BT.31 (Pl. 6, Fig. 7): several dorsal valves illustrating the structure and development of the lophophore. Aspects of internal ventral parts of the animal. ZMB���Bra 2472 (Pl. 6, Fig. 6): ventral valve of a female specimen, preserved in ethanol, observed under binocular and showing the gonad with spherical female gametes. Etymology. The species name is derived from ���Mawali��� the name of the shipwreck where it has been found for the first time. Type locality. The ���Mawali��� shipwreck (1��26���46.74���N / 125��13���32.16���E), Lembeh Strait, North Sulawesi Province, Indonesia. Additional material. The material studied is composed of two types of samples. The first type is dried material collected either from sieved sediment or from specimens scraped off the walls of the hold or living specimens attached on oyster shells. The second type consists of oyster shells with living specimens directly placed in ethanol by the collector. The dried material represents 285 specimens whereas the material preserved in ethanol consists of 206 specimens (both juveniles and adults). Note that many specimens in this latter sample were already dead but still attached to their support and that truly living specimens when collected were less abundant. Diagnosis. Small-sized Thecidellina species. Elongated drop-like ventral valve with striated, triangular, flat planodeltidium. Hemispondylium arising from posterior part of the valve, connected to the ventral valve floor for only a short part of its length, with long and slender prongs acutely pointed reaching the level of the cyrtomatodont teeth. Ventral valve floor without median ridge, very coarsely tuberculate except in the deep ovate gonad pits, which have a smooth surface. Dorsal valve lid-like. Brachial bridge and intrabrachial ridge firmly connected by wide posterior outgrowths. Calcitic pole, thick, attached to dorsal valve floor and connected with anterior expansions to the posterior parts of the brachial bridge in fully adult specimens. Median septum high, thin, with acute tip, ventrally convex, tuberculate-spinous on ventral surface. Brachial cavities or brood pouches covered with thin, fragile irregular canopying spicules that become fused together through ontogeny but leaving always a lot of randomly disposed small cavities at maturity. Mendiagnosia. Thecidellina spesies berukuran kecil. Memanjang katup ventral tetes berbentuk dengan lurik, segitiga planodeltidium. Hemispondylium timbul dari bagian posterior katup, tak lama terhubung ke lantai katup ventral, berbentuk garpu dengan dua pisau panjang dan ramping mencapai tingkat gigi cyrtomatodont. Lantai katup ventral, tanpa ridge, sangat kasar tuberculate kecuali gonads lubang-lubang dengan permukaan halus. Katup dorsal seperti penutup kaleng. Jembatan brakialis dan punggung peribrachial tegas tersambung dengan posterior persimpangan piring-piring. Calcitic pole, tebal, bergabung ke lantai katup ventral. Dalam spesimen sepenuhnya dewasa, bagian depan dari calcitic pole juga dihubungkan oleh ekspansinya ke bagian posterior jembatan brakialis. Median septum tinggi, dengan ujung akut, ventally cembung, dengan tuberkel kecil di permukaan ventral. Brakialis parit-parit (atau merenung kantong) ditutupi dengan tipis dan rapuh spikulae sekering bersama-sama melalui ontogeni untuk berbentuk satu cakupan tetapi selalu meninggalkan rongga-rongga kecil dengan bentukbentuk sangat variabel. Description. External shell characters. Relatively small-sized, whitish thecideide (Table 1), with endopunctate shell, slightly longer than wide in adult stage but often as long as wide in juvenile stages of growth (Pl. 1, Figs. 1a, 2a; Pl. 2, Figs. 1a, 2a). The external surface of the shell is marked by many growth lines with variable thickness. The shell is strongly ventri-biconvex, with the maximum thickness situated at mid-length or slightly backward of this for the ventral valve. The maximum thickness for the dorsal valve corresponds to the position of the protegulum (Pl. 1, Figs. 1c, 2d). The lateral commissure is variable being dorsally concave (Pl. 2, Fig. 2c), rectimarginate (Pl. 2, Fig. 1d) or slightly sinusoidal (Pl. 1, Figs. 1c, 2d). The anterior commissure is rectimarginate (Pl. 1, Fig. 2f). The deep ventral valve has an ellipsoid to hemispherical outline in lateral profile (Pl. 1, Figs. 1e, 2d) and a very faint sulcus is sometimes developed in the middle of the anterior part of the valve (Pl. 1, Fig. 2e���f). The ventral valve is cemented to the substrate by its posterior part and often by a large portion of its ventral side (Pl. 1, Fig. 2e). The adult shell is often lifted from the substrate anteriorly (Pl. 1, Fig. 1e), its anterior part tending to be always more elevated than its posterior part. Such a development of the ventral valve is common in thecideide brachiopods as already illustrated by Pajaud (1970, p. 219, fig. 130A). The interarea is a planodeltidium (Logan & Baker 2013) with an isosceles triangular outline, and a flat (Pl. 1, Fig. 2d) or slightly concave (Pl. 1, Fig. 1d���e) surface. The interarea represents 22���31% of the length of the shell (Table 1). Due to variable habitat conditions the planodeltidium can be curved to the left or the right sides (Pl. 2, Figs. 1a, 3). The surface of the interarea is striated with parallel growth lines. The hinge line is straight without a notch. The interarea and the triangular posterior part of the dorsal valve form a sharp angle at the level of the hinge line (Pl. 1, Figs. 1a���b, 2a���b) but the value of this angle is highly variable (Pl. 1, Fig. 2d; Pl. 2, Fig. 1c). The dorsal valve, much smaller than the ventral valve, has a lid-shaped outline, subcircular to elongate oval and it is sometimes slightly emarginate (if a faint sulcus is developed on the anterior part of the ventral valve). The weak convexity of this valve culminates posteriorly with the protegulum. The surface of the dorsal valve is smooth except for the development of irregular growth lines. The protegulum is circular, always very distinct and its external surface is covered with numerous pustules (Pl. 1, Figs. 1c, 2c; Pl. 2, Fig. 2f). The posterior part of the dorsal valve is obtusely triangular with a flat striated surface. Most of the morphological characters increase regularly and proportionally during growth as indicated by the linear relations observed between width, length, length of dorsal valve and width of the hinge (Text-Fig. 3). However, the thickness of the shell follows a sigmoid development. This is due to the fact that juveniles are extremely flat and the thickness is increasing very slowly at the beginning of growth. When the shell length reaches 2 mm, the thickness suddenly increases very rapidly. This type of growth is commonly observed in the development of Thecidellina species (Text-Fig. 3). Internal shell characters. The ventral valve has an internal subcircular or sometimes subcordiform outline (Pl. 3, Fig. 2a). A ventrally concave, narrow, smooth peripheral rim is developed along the commissure (Pl. 3, Fig. 2a, 2c���d). Along the internal side of this rim, a row of regular tubercles is developed (Pl. 3, Fig. 2a���c). The ventral valve floor, without a differentiated median ridge, is strongly tuberculate (with tubercles of regular size) except inside the two very deep oval pits where the gonads are situated (Pl. 3, Fig. 2a, 2c, 2g). The gonad pits are relatively large in this species, their length being frequently up to 40% of internal valve length (Pl. 3, Fig. 2a). The endopunctation is clearly visible between the tubercles of the valve floor. The hemispondylium is made of two parallel, narrow plates with long pointed curved prongs attached to the posterior part of the valve and also to the valve floor for a very short distance (Pl. 3, Fig. 2e). The delthyrial cavity extends under the planodeltidium and has a wide dorsal ridge forming, in anterior view, the form of a letter ���M��� (Pl. 3, Fig. 2e). Large, oval kidney-shaped lateral adductor muscle scars are developed on either side of the hemispondylium and teeth (Pl. 3, Fig. 2f). The cyrtomatodont, robust and short teeth are covered with secondary shell material and are obtusely pointed (Pl. 3, Fig. 2a, 2d���e). The dorsal valve is slightly convex, subcircular with a cordiform anterior outline. A flat, relatively smooth flange is externally visible along the commissure. The external side of the peripheral ridge is ornamented with thick tubercles and its crest shows regular tubercles till its junction with the brachial bridge (Pl. 3, Fig. 1h). A clearly defined peripheral rim is not present. In lateral profile the internal structures are raised towards the posterior part (Pl. 3, Fig. 1h). The angle formed between the anterior limit of the valve, the posterior part of the peripheral ridge and the top of the brachial bridge has a value of 17.5 degrees (Pl. 3, Fig. 1j). The median septum is straight, wide anteriorly, tapering posteriorly, and very thin in its posterior part. In lateral view the median septum is ventrally convex. The crest of the septum is covered with one row of tubercles decreasing in size from the anterior base to the tip (Pl. 3, Fig. 1g). The intrabrachial ridge is clearly defined and is roughly tuberculate (Pl. 3, Fig. 1h). Marsupial orifices are situated in the posterior part of the intrabrachial ridge, on either side of the septal tip (Pl. 3, Fig. 1a, 1g). These orifices are small sub-circular holes (Pl. 5, Fig. 2a, 2e) confined by an extremely thin and fragile calcitic rim on their ventral side. For this reason, they appear often broken as an open circle (Pl. 3, Fig. 1a���b; Pl. 5, Figs. 5a, 8a���b). The lophophore groove is relatively deep and wide (Pl. 3, Fig. 1a). In the posterior part of the valve, two large lateral visceral gaps are situated between the intrabrachial ridge and the brachial bridge (���vg���, Pl. 3, Fig. 1h). The posterior visceral gap between the brachial bridge and the intrabrachial ridge is relatively narrow (Pl. 3, Fig. 1a). The brachial bridge has a finely denticulated ventral crest (Pl. 3, Fig. 1j���k). The internal side of the brachial bridge exhibits numerous lophophore muscle scars. Clearly defined, posterior wide outgrowths firmly connect the inner side of the brachial bridge and the posterior side of the intrabrachial ridge in adult specimens (���og���, Pl. 3, Fig. 1e). The cardinal process is short, massive, trilobate and dorsally curved (���cpr���, Pl. 3, Fig. 1a, 1j���l; Pl. 5, Figs. 6, 7, 8a). At the posterior part of the cardinal process, the diductor muscle scars have a triangular outline (���did���, Pl. 3, Fig. 1k). Inner socket ridges are thickened and quite strong (Pl. 3, Fig. 1i���j). Outer socket ridges are flat and thin. The sockets are moderately deep. Placed on either side of the base of the inner socket ridges, the lateral adductor muscle scars are widely developed (Pl. 3, Fig. 1j���l). The anterior side of the calcitic pole is narrow and has two wide lateral expansions (���Exp���, Pl. 3, Fig. 1f). These expansions are fused with the brachial bridge at maturity (Pl. 5, Figs. 6, 7, 8a���b). Seen from behind, the calcitic pole is thick, fused with the dorsal valve floor at the anterior part of the low but long and posteriorly pointed spur of the cardinal process (Pl. 3, Fig. 1k���l). The brachial cavities (brood pouches in female specimens) are deep and covered with spiculate canopies on their half anterior part. Canopying spicules are thin, irregular in shape and interconnected, producing a quite strong cover protecting the embryos. Several small holes occur in the canopy structure (Pl. 3, Fig. 1a). These holes are variable in number and size. The posterior parts of brachial cavities remain uncovered. Canopy spicules have a fine granulated structure ornamenting their ventral side (Pl. 3, Fig. 1a���b). This granulation, observed with high SEM magnification, consists of pointed subpyramidal prisms with acute tips (Pl. 3, Fig. 1c���d). The canopies are built with spicules situated on the external margins of the brachial cavities (Pl. 3, Fig. 1a). However, spicules situated on the internal margins can also participate in the erection of canopies (Pl. 5, Figs. 3, 5a, 7). This latest development of the spicules could be related to the growth stage and appearing only in fully mature specimens. Shell ontogeny. At its youngest stage of growth, the ventral valve presents all the characteristics of its adult morphology: the flat triangular planodeltidium, the hinge structure, the rough internal valve floor and the bifid hemispondylium. In the contrast, the dorsal valve undergoes a great transformation giving rise to numerous morphological characters of generic and specific value. The youngest dorsal valve stage of growth found in our material (Pl. 4, Fig. 1a���d) shows strong socket ridges and a developed, bilobed, cardinal process. A clear peripheral rim is visible in the posterolateral parts of the valve and the peribrachial ridge is delimited by a dozen small dispersed tubercles. The brachial bridge is already completely developed. Two separate spikes are erected in the centre of the valve floor: they are flat laterally but their tips are pointed and oriented backwards. The adductor muscle scars are also clearly visible (Pl. 4, Fig. 1d). The next stage of growth (Pl. 4, Fig. 2a���c) shows a cardinal process with a smooth median spur. The lateral lobes of the cardinal process are covered with secondary fibers. All the muscle scars are situated on a triangular platform at the posterior end of the cardinal process. Note also that the structure of the hemispondylium is completely developed (Pl. 4, Fig. 2a���b). It is made of two separate long, pointed claws which for only a short part of their length are attached to the ventral valve floor. The peribrachial ridge is more developed with an increasing number of tubercles. The two spikes in the center of the dorsal valve floor are now connected with the emergent septum which is still rather short. The intrabrachial ridge begins its development by the appearance of lateral extension(s) growing from the base of the original spikes to the lateral parts of the shell floor (Pl. 4, Figs. 2a, 2c, 3b). The development of these posterior parts of the intrabrachial ridge is better illustrated in Pl. 4, Fig. 3a���c where symmetrical progressive outgrowths are visible. The central spikes become longer increasing in length posteriorly. The median septum is slightly higher and more developed. At this stage of development two small prominent outgrowths produced just at the base of the inner side of the brachial bridge first appear; they increase in length anteriorly (Pl. 4, Fig. 3a, 3e). These outgrowths later connect the brachial bridge and the intrabrachial ridge together. In Thecidellina mawaliana sp. nov. they appear very early in ontogeny when compared with other species of Thecidellina. Nevertheless, in some cases (for unknown reasons) the development of these outgrowths could be variable during ontogeny appearing later in development, as seen in one illustrated specimen (Pl. 5, Fig. 1a), or they remain very small. In this case, the two spikes at the beginning of ontogeny that are now at the tip of the posterior part of the median septum, continue to grow posteriorly to the peribrachial ridge and become thickened (Pl. 5, Fig. 1e). Finally, they join the small outgrowths from the brachial bridge connecting it to the peribrachial ridge (pl. 5, Fig. 3). The intrabrachial ridge is under construction with the emergence of small tubercles that are quite discrete at the beginning (Pl. 4, Fig. 3a���d) but which increase in thickness and height progressively (Pl. 4, Fig. 4a���c). No calcitic pole is developed at this stage of growth. Afterwards the median septum increases in length and joins the anterior part of the peribrachial ridge where it becomes clearly wider (Pl. 5, Fig. 1a���b, 1e). At this stage, this median septum remains relatively thin but its height reaches its maximum development at its posterior end where it is fused with the intrabrachial ridge (Pl. 5, Fig. 1c). A thin, lamellar calcitic pole is produced at this stage of growth. It develops in length more in its posterior part than in its anterior part (Pl. 5, Fig. 1f���g). The tubercles which initiated the construction of the peribrachial ridge are now fused together building the brachial cavities. Canopying spicules emerge progressively. This process begins in the posterior portion of the peribrachial ridge (Pl. 5, Fig. 1b���c, 1e), continues through its lateral parts (Pl. 5, Fig. 4) and is finally achieved in its anterior part (Pl. 5, Figs. 5a, 6���7, 8a). The next step is the interconnection between the base of the brachial bridge and the posterior side of the peribrachial ridge. This interconnection is completed generally by the development of the posterior prominent outgrowths (Pl. 5, Fig. 2a, 2e). These outgrowths are also connected now with the anterior lateral expansions of the calcitic pole (Pl. 5, Fig. 2e). From these interconnections between brachial bridge and intrabrachial ridge appear three visceral gaps: two elongated lateral gaps and one posterior visceral gap. All these c, Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on pages 485-492, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Pajaud, D. (1970) Monographie des Thecidees. Memoires de la Societe geologique de France, Nouvelle Serie, 49 (112), 1 - 349.","Logan, A. & Baker, P. (2013) The development and shell microstructure of the pseudodeltidium and interarea in thecideide brachiopods. Palaeontology, 56, 433 - 455. https: // doi. org / 10.1111 / pala. 12001"]}

SUBPHYLUM RHYNCHONELLIFORMEA WILLIAMS ET AL., 1996 CLASS RHYNCHONELLATA WILLIAMS ET AL., 1996, Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 485, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832

Thecidellina mawaliana sp. nov. Table 1; Text-Figs. 2–4; Pls. 1–6 Holotype: RBINS–BT.9 (Pl. 1, Fig. 2a–g, Pl. 5, Fig. 4): articulated adult specimen. Paratypes: RBINS–BT.10 (Pl. 1, Fig. 1a–f): articulated adult specimen. RBINS–BT.11 (Pl. 3, Fig. 1a–n): adult dorsal valve with complete development of the canopies on brachial cavities. RBINS–BT.12 (Pl. 2, Fig. 1a–g): small articulated specimen with brachial cavities still without canopying spicules and with a planodeltidium curving to right side. RBINS–BT.13 (Pl. 2, Fig. 2a–g, Pl. 4, Fig. 2a–c): juvenile articulated specimen. RBINS– BT.14 (Pl. 2, Fig. 3): small articulated specimen with its planodeltidium curving to the left side. RBINS–BT.15 (Pl. 3, Fig. 2a–f): adult ventral valve with teeth and intact hemispondylium. RBINS–BT.16 (Pl. 4, Fig. 1a–d): a very early juvenile dorsal valve. RBINS–BT.17 (Pl. 4, Fig. 3a–f): juvenile dorsal valve with a peribrachial ridge beginning its development and with appearing posterior outgrowths from the brachial bridge. RBINS–BT.18 (Pl. 4, Fig. 4a–c): juvenile dorsal valve with tubercles beginning the building of the brachial cavities. RBINS–BT.19 (Pl. 5, Fig. 1a–g): dorsal valve with developed median septum, first appearance of calcitic pole and continuing the development of brachial cavities. RBINS–BT.20 (Pl. 5, Fig. 2a–e): dorsal valve with brachial bridge and peribrachial ridge connected. RBINS–BT.21 (Pl. 5, Fig. 3): adult dorsal valve with brachial cavities with incomplete canopies. RBINS–BT.22 (Pl. 5, Fig. 6), RBINS–BT.23 (Pl. 5, Fig. 7), RBINS–BT.24 (Pl. 5, Fig. 5a–b), and RBINS–BT.25 (Pl. 5, Fig. 8a–b): several adult dorsal valves illustrating the variation of the canopies covering the brachial cavities. RBINS–BT.26 (Pl. 6, Fig. 1), RBINS–BT.27 (Pl. 6, Fig. 2), RBINS–BT.28 (Pl. 6, Fig. 3), RBINS–BT.29 (Pl. 6, Fig. 4), RBINS–BT.30 (Pl. 6, Fig. 5), and RBINS–BT.31 (Pl. 6, Fig. 7): several dorsal valves illustrating the structure and development of the lophophore. Aspects of internal ventral parts of the animal. ZMB–Bra 2472 (Pl. 6, Fig. 6): ventral valve of a female specimen, preserved in ethanol, observed under binocular and showing the gonad with spherical female gametes. Etymology. The species name is derived from “Mawali” the name of the shipwreck where it has been found for the first time. Type locality. The “Mawali” shipwreck (1°26’46.74”N / 125°13’32.16”E), Lembeh Strait, North Sulawesi Province, Indonesia. Additional material. The material studied is composed of two types of samples. The first type is dried material collected either from sieved sediment or from specimens scraped off the walls of the hold or living specimens attached on oyster shells. The second type consists of oyster shells with living specimens directly placed in ethanol by the collector. The dried material represents 285 specimens whereas the material preserved in ethanol consists of 206 specimens (both juveniles and adults). Note that many specimens in this latter sample were already dead but still attached to their support and that truly living specimens when collected were less abundant. Diagnosis. Small-sized Thecidellina species. Elongated drop-like ventral valve with striated, triangular, flat planodeltidium. Hemispondylium arising from posterior part of the valve, connected to the ventral valve floor for only a short part of its length, with long and slender prongs acutely pointed reaching the level of the cyrtomatodont teeth. Ventral valve floor without median ridge, very coarsely tuberculate except in the deep ovate gonad pits, which have a smooth surface. Dorsal valve lid-like. Brachial bridge and intrabrachial ridge firmly connected by wide posterior outgrowths. Calcitic pole, thick, attached to dorsal valve floor and connected with anterior expansions to the posterior parts of the brachial bridge in fully adult specimens. Median septum high, thin, with acute tip, ventrally convex, tuberculate-spinous on ventral surface. Brachial cavities or brood pouches covered with thin, fragile irregular canopying spicules that become fused together through ontogeny but leaving always a lot of randomly disposed small cavities at maturity. Mendiagnosia. Thecidellina spesies berukuran kecil. Memanjang katup ventral tetes berbentuk dengan lurik, segitiga planodeltidium. Hemispondylium timbul dari bagian posterior katup, tak lama terhubung ke lantai katup ventral, berbentuk garpu dengan dua pisau panjang dan ramping mencapai tingkat gigi cyrtomatodont. Lantai katup ventral, tanpa ridge, sangat kasar tuberculate kecuali gonads lubang-lubang dengan permukaan halus. Katup dorsal seperti penutup kaleng. Jembatan brakialis dan punggung peribrachial tegas tersambung dengan posterior persimpangan piring-piring. Calcitic pole, tebal, bergabung ke lantai katup ventral. Dalam spesimen sepenuhnya dewasa, bagian depan dari calcitic pole juga dihubungkan oleh ekspansinya ke bagian posterior jembatan brakialis. Median septum tinggi, dengan ujung akut, ventally cembung, dengan tuberkel kecil di permukaan ventral. Brakialis parit-parit (atau merenung kantong) ditutupi dengan tipis dan rapuh spikulae sekering bersama-sama melalui ontogeni untuk berbentuk satu cakupan tetapi selalu meninggalkan rongga-rongga kecil dengan bentukbentuk sangat variabel. Description. External shell characters. Relatively small-sized, whitish thecideide (Table 1), with endopunctate shell, slightly longer than wide in adult stage but often as long as wide in juvenile stages of growth (Pl. 1, Figs. 1a, 2a; Pl. 2, Figs. 1a, 2a). The external surface of the shell is marked by many growth lines with variable thickness. The shell is strongly ventri-biconvex, with the maximum thickness situated at mid-length or slightly backward of this for the ventral valve. The maximum thickness for the dorsal valve corresponds to the position of the protegulum (Pl. 1, Figs. 1c, 2d). The lateral commissure is variable being dorsally concave (Pl. 2, Fig. 2c), rectimarginate (Pl. 2, Fig. 1d) or slightly sinusoidal (Pl. 1, Figs. 1c, 2d). The anterior commissure is rectimarginate (Pl. 1, Fig. 2f). The deep ventral valve has an ellipsoid to hemispherical outline in lateral profile (Pl. 1, Figs. 1e, 2d) and a very faint sulcus is sometimes developed in the middle of the anterior part of the valve (Pl. 1, Fig. 2e–f). The ventral valve is cemented to the substrate by its posterior part and often by a large portion of its ventral side (Pl. 1, Fig. 2e). The adult shell is often lifted from the substrate anteriorly (Pl. 1, Fig. 1e), its anterior part tending to be always more elevated than its posterior part. Such a development of the ventral valve is common in thecideide brachiopods as already illustrated by Pajaud (1970, p. 219, fig. 130A). The interarea is a planodeltidium (Logan & Baker 2013) with an isosceles triangular outline, and a flat (Pl. 1, Fig. 2d) or slightly concave (Pl. 1, Fig. 1d–e) surface. The interarea represents 22–31% of the length of the shell (Table 1). Due to variable habitat conditions the planodeltidium can be curved to the left or the right sides (Pl. 2, Figs. 1a, 3). The surface of the interarea is striated with parallel growth lines. The hinge line is straight without a notch. The interarea and the triangular posterior part of the dorsal valve form a sharp angle at the level of the hinge line (Pl. 1, Figs. 1a–b, 2a–b) but the value of this angle is highly variable (Pl. 1, Fig. 2d; Pl. 2, Fig. 1c). The dorsal valve, much smaller than the ventral valve, has a lid-shaped outline, subcircular to elongate oval and it is sometimes slightly emarginate (if a faint sulcus is developed on the anterior part of the ventral valve). The weak convexity of this valve culminates posteriorly with the protegulum. The surface of the dorsal valve is smooth except for the development of irregular growth lines. The protegulum is circular, always very distinct and its external surface is covered with numerous pustules (Pl. 1, Figs. 1c, 2c; Pl. 2, Fig. 2f). The posterior part of the dorsal valve is obtusely triangular with a flat striated surface. Most of the morphological characters increase regularly and proportionally during growth as indicated by the linear relations observed between width, length, length of dorsal valve and width of the hinge (Text-Fig. 3). However, the thickness of the shell follows a sigmoid development. This is due to the fact that juveniles are extremely flat and the thickness is increasing very slowly at the beginning of growth. When the shell length reaches 2 mm, the thickness suddenly increases very rapidly. This type of growth is commonly observed in the development of Thecidellina species (Text-Fig. 3). Internal shell characters. The ventral valve has an internal subcircular or sometimes subcordiform outline (Pl. 3, Fig. 2a). A ventrally concave, narrow, smooth peripheral rim is developed along the commissure (Pl. 3, Fig. 2a, 2c–d). Along the internal side of this rim, a row of regular tubercles is developed (Pl. 3, Fig. 2a–c). The ventral valve floor, without a differentiated median ridge, is strongly tuberculate (with tubercles of regular size) except inside the two very deep oval pits where the gonads are situated (Pl. 3, Fig. 2a, 2c, 2g). The gonad pits are relatively large in this species, their length being frequently up to 40% of internal valve length (Pl. 3, Fig. 2a). The endopunctation is clearly visible between the tubercles of the valve floor. The hemispondylium is made of two parallel, narrow plates with long pointed curved prongs attached to the posterior part of the valve and also to the valve floor for a very short distance (Pl. 3, Fig. 2e). The delthyrial cavity extends under the planodeltidium and has a wide dorsal ridge forming, in anterior view, the form of a letter “M” (Pl. 3, Fig. 2e). Large, oval kidney-shaped lateral adductor muscle scars are developed on either side of the hemispondylium and teeth (Pl. 3, Fig. 2f). The cyrtomatodont, robust and short teeth are covered with secondary shell material and are obtusely pointed (Pl. 3, Fig. 2a, 2d–e). The dorsal valve is slightly convex, subcircular with a cordiform anterior outline. A flat, relatively smooth flange is externally visible along the commissure. The external side of the peripheral ridge is ornamented with thick tubercles and its crest shows regular tubercles till its junction with the brachial bridge (Pl. 3, Fig. 1h). A clearly defined peripheral rim is not present. In lateral profile the internal structures are raised towards the posterior part (Pl. 3, Fig. 1h). The angle formed between the anterior limit of the valve, the posterior part of the peripheral ridge and the top of the brachial bridge has a value of 17.5 degrees (Pl. 3, Fig. 1j). The median septum is straight, wide anteriorly, tapering posteriorly, and very thin in its posterior part. In lateral view the median septum is ventrally convex. The crest of the septum is covered with one row of tubercles decreasing in size from the anterior base to the tip (Pl. 3, Fig. 1g). The intrabrachial ridge is clearly defined and is roughly tuberculate (Pl. 3, Fig. 1h). Marsupial orifices are situated in the posterior part of the intrabrachial ridge, on either side of the septal tip (Pl. 3, Fig. 1a, 1g). These orifices are small sub-circular holes (Pl. 5, Fig. 2a, 2e) confined by an extremely thin and fragile calcitic rim on their ventral side. For this reason, they appear often broken as an open circle (Pl. 3, Fig. 1a–b; Pl. 5, Figs. 5a, 8a–b). The lophophore groove is relatively deep and wide (Pl. 3, Fig. 1a). In the posterior part of the valve, two large lateral visceral gaps are situated between the intrabrachial ridge and the brachial bridge (“vg”, Pl. 3, Fig. 1h). The posterior visceral gap between the brachial bridge and the intrabrachial ridge is relatively narrow (Pl. 3, Fig. 1a). The brachial bridge has a finely denticulated ventral crest (Pl. 3, Fig. 1j–k). The internal side of the brachial bridge exhibits numerous lophophore muscle scars. Clearly defined, posterior wide outgrowths firmly connect the inner side of the brachial bridge and the posterior side of the intrabrachial ridge in adult specimens (“og”, Pl. 3, Fig. 1e). The cardinal process is short, massive, trilobate and dorsally curved (“cpr”, Pl. 3, Fig. 1a, 1j–l; Pl. 5, Figs. 6, 7, 8a). At the posterior part of the cardinal process, the diductor muscle scars have a triangular outline (“did”, Pl. 3, Fig. 1k). Inner socket ridges are thickened and quite strong (Pl. 3, Fig. 1i–j). Outer socket ridges are flat and thin. The sockets are moderately deep. Placed on either side of the base of the inner socket ridges, the lateral adductor muscle scars are widely developed (Pl. 3, Fig. 1j–l). The anterior side of the calcitic pole is narrow and has two wide lateral expansions (“Exp”, Pl. 3, Fig. 1f). These expansions are fused with the brachial bridge at maturity (Pl. 5, Figs. 6, 7, 8a–b). Seen from behind, the calcitic pole is thick, fused with the dorsal valve floor at the anterior part of the low but long and posteriorly pointed spur of the cardinal process (Pl. 3, Fig. 1k–l). The brachial cavities (brood pouches in female specimens) are deep and covered with spiculate canopies on their half anterior part. Canopying spicules are thin, irregular in shape and interconnected, producing a quite strong cover protecting the embryos. Several small holes occur in the canopy structure (Pl. 3, Fig. 1a). These holes are variable in number and size. The posterior parts of brachial cavities remain uncovered. Canopy spicules have a fine granulated structure ornamenting their ventral side (Pl. 3, Fig. 1a–b). This granulation, observed with high SEM magnification, consists of pointed subpyramidal prisms with acute tips (Pl. 3, Fig. 1c–d). The canopies are built with spicules situated on the external margins of the brachial cavities (Pl. 3, Fig. 1a). However, spicules situated on the internal margins can also participate in the erection of canopies (Pl. 5, Figs. 3, 5a, 7). This latest development of the spicules could be related to the growth stage and appearing only in fully mature specimens. Shell ontogeny. At its youngest stage of growth, the ventral valve presents all the characteristics of its adult morphology: the flat triangular planodeltidium, the hinge structure, the rough internal valve floor and the bifid hemispondylium. In the contrast, the dorsal valve undergoes a great transformation giving rise to numerous morphological characters of generic and specific value. The youngest dorsal valve stage of growth found in our material (Pl. 4, Fig. 1a–d) shows strong socket ridges and a developed, bilobed, cardinal process. A clear peripheral rim is visible in the posterolateral parts of the valve and the peribrachial ridge is delimited by a dozen small dispersed tubercles. The brachial bridge is already completely developed. Two separate spikes are erected in the centre of the valve floor: they are flat laterally but their tips are pointed and oriented backwards. The adductor muscle scars are also clearly visible (Pl. 4, Fig. 1d). The next stage of growth (Pl. 4, Fig. 2a–c) shows a cardinal process with a smooth median spur. The lateral lobes of the cardinal process are covered with secondary fibers. All the muscle scars are situated on a triangular platform at the posterior end of the cardinal process. Note also that the structure of the hemispondylium is completely developed (Pl. 4, Fig. 2a–b). It is made of two separate long, pointed claws which for only a short part of their length are attached to the ventral valve floor. The peribrachial ridge is more developed with an increasing number of tubercles. The two spikes in the center of the dorsal valve floor are now connected with the emergent septum which is still rather short. The intrabrachial ridge begins its development by the appearance of lateral extension(s) growing from the base of the original spikes to the lateral parts of the shell floor (Pl. 4, Figs. 2a, 2c, 3b). The development of these posterior parts of the intrabrachial ridge is better illustrated in Pl. 4, Fig. 3a–c where symmetrical progressive outgrowths are visible. The central spikes become longer increasing in length posteriorly. The median septum is slightly higher and more developed. At this stage of development two small prominent outgrowths produced just at the base of the inner side of the brachial bridge first appear; they increase in length anteriorly (Pl. 4, Fig. 3a, 3e). These outgrowths later connect the brachial bridge and the intrabrachial ridge together. In Thecidellina mawaliana sp. nov. they appear very early in ontogeny when compared with other species of Thecidellina. Nevertheless, in some cases (for unknown reasons) the development of these outgrowths could be variable during ontogeny appearing later in development, as seen in one illustrated specimen (Pl. 5, Fig. 1a), or they remain very small. In this case, the two spikes at the beginning of ontogeny that are now at the tip of the posterior part of the median septum, continue to grow posteriorly to the peribrachial ridge and become thickened (Pl. 5, Fig. 1e). Finally, they join the small outgrowths from the brachial bridge connecting it to the peribrachial ridge (pl. 5, Fig. 3). The intrabrachial ridge is under construction with the emergence of small tubercles that are quite discrete at the beginning (Pl. 4, Fig. 3a–d) but which increase in thickness and height progressively (Pl. 4, Fig. 4a–c). No calcitic pole is developed at this stage of growth. Afterwards the median septum increases in length and joins the anterior part of the peribrachial ridge where it becomes clearly wider (Pl. 5, Fig. 1a–b, 1e). At this stage, this median septum remains relatively thin but its height reaches its maximum development at its posterior end where it is fused with the intrabrachial ridge (Pl. 5, Fig. 1c). A thin, lamellar calcitic pole is produced at this stage of growth. It develops in length more in its posterior part than in its anterior part (Pl. 5, Fig. 1f–g). The tubercles which initiated the construction of the peribrachial ridge are now fused together building the brachial cavities. Canopying spicules emerge progressively. This process begins in the posterior portion of the peribrachial ridge (Pl. 5, Fig. 1b–c, 1e), continues through its lateral parts (Pl. 5, Fig. 4) and is finally achieved in its anterior part (Pl. 5, Figs. 5a, 6–7, 8a). The next step is the interconnection between the base of the brachial bridge and the posterior side of the peribrachial ridge. This interconnection is completed generally by the development of the posterior prominent outgrowths (Pl. 5, Fig. 2a, 2e). These outgrowths are also connected now with the anterior lateral expansions of the calcitic pole (Pl. 5, Fig. 2e). From these interconnections between brachial bridge and intrabrachial ridge appear three visceral gaps: two elongated lateral gaps and one posterior visceral gap. All these c

Genus Minutella Hoffmann & L��ter, 2010 Type species. Minutella tristani Hoffmann & L��ter, 2010., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 495, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Hoffmann, J. & Luter, C. (2010) Shell development in thecidellinine brachiopods with description of a new Recent genus. Special Papers in Palaeontology, 84, 137 - 160."]}

Genus Ospreyella L��ter & W��rheide in L��ter et al., 2003 Type species. Ospreyella depressa L��ter in L��ter et al., 2003., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 496, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Luter, C., Worheide, G. & Reitner, J. (2003) A new thecideid genus and species (Brachiopoda, Recent) from submarine caves of Osprey Reef (Queensland Plateau, Coral Sea, Australia). Journal of Natural History, 37, 1423 - 1432. https: // doi. org / 10.1080 / 00222930110120971"]}

Comparison with Thecidellina blochmanni Dall, 1920 1920 Thecidellina blochmanni: Dall, p. 283. 1939 Thecidellina blochmanni: Helmcke, pp. 216���233, fig. 231. 1965 Thecidellina blochmanni: Elliott, fig. 742.3a���b. 1973 Thecidellina blochmanni: Cooper, p. 8, pl. 8, figs. 27���30. 1985 Thecidellina blochmanni: Zezina, p. 208. ? 1987 Thecidellina blochmanni: D���Hondt, p. 41. 2003 Thecidellina blochmanni: Lee & Robinson, p. 354. pp 2013 Thecidellina blochmanni: Logan & Bitner, fig. 4F, non fig. 4A���E, 4G���L. 2015 Thecidellina blochmanni: Logan et al., pp. 229���232, fig. 3A���O. This species from Christmas Island (Australia, Indian Ocean) has a wider median septum in dorsal valve with stronger imprints of lophophore scars on its lateral sides than T. mawaliana sp. nov. In T. blochmanni, the brachial cavities are deeper and covered with very strong, thick angular spicules which were described ���massive bouffantlike calcitic aggregations��� by Logan et al. (2015, p. 230). The thin, irregular, fused spicules of T. mawaliana sp. nov. are quite different. Remarks. ? T. blochmanni in D���Hondt (1987): the species is just cited without description and without illustration. This material must be revisited., Published as part of Simon, Eric, L��ter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on pages 493-494, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Dall, W. H. (1920) Annotated list of the Recent Brachiopoda in the collection of the United States National Museum, with description of thirty-three new forms. Proceedings of the United States National Museum, 57 (2314), 261 - 377. https: // doi. org / 10.5479 / si. 00963801.57 - 2314.261","Logan, A., Hoffmann, J. & Luter, C. (2015) Checklist of Recent thecideoid brachiopods from the Indian Ocean and Red Sea, with a description of a new species of Thecidellina from Europa Island and a re-description of T. blochmanni Dall from Christmas Island. Zootaxa, 4013 (2), 225 - 234. https: // doi. org / 10.11646 / zootaxa. 4013.2.4"]}

Comparison with Thecidellina japonica (Hayasaka, 1938) 1938 Thecidellella japonica: Hayasaka, pp. 11–12; figs. 1–2. 1940 Thecidellella japonica: Hatai, pp. 192–193, fig. 19. 1985 Thecidellina japonica: Zezina, p. 208. In T. japonica, the ventral valve floor is smooth with a very characteristic median ridge made of three longitudinal series of irregular small tubercles. The dorsal valve has brachial cavities without covering canopies. In T. mawaliana nov. sp. no median ridge is developed (undifferentiated) and the median floor is regularly covered with tubercles., Published as part of Simon, Eric, Lüter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on page 494, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Hayasaka, L. (1938) A new Neotreme genus of Brachiopod from Japan. The Fenus, 8, 9 - 13."]}

Minutella cf. minuta (Cooper, 1981) (Lembeh form) Text-Fig. 2; Pl. 7, Figs 1a–g, 2 ? 1981 Thecidellina minuta: Cooper, p. 62; pl. 6, figs. 27–40. ? 2013 Minutella cf. minuta: Simon & Hoffmann, p. 405; pl. 1, figs. 1–5, pl. 2, figs. 1–8. ? 2016 Minutella cf. minuta (Cooper, 1981): Simon et al., p. 5. Material. Available specimens were collected in the shipwreck from sieved sediment or attached on oyster shells and preserved in ethanol. Specimens in ethanol, although numerous (111), are reserved for biological studies. The dried or dead specimens of this species are represented in this station with a lower quality level of preservation. The dried material is represented by 15 articulated shells, 19 dorsal and 6 ventral valves. Description. The shell is elongate, drop-like with a pointed umbo in outline, strongly biconvex in lateral and anterior views. In posterior view, the bottom of the shell (Pl. 7, Fig. 2) is flat when visible (the shell is often attached by its posterior side). The anterior commissure is rectimarginate whereas the lateral commissure is straight or slightly concave dorsally. The ventral valve has a smooth external surface with many fine growth lines. The transversely striated triangular interarea (Pl. 7, Fig. 1g) is flat or presents a very unlikely concavity and flanks a wide, dorsally convex rugideltidium (sensu Logan and Baker 2013). The smooth peripheral rim is relatively wide and the peripheral ridge has one or two rows of small tubercles (Pl. 7, Fig. 1g). The ventral valve floor is smooth. The hemispondylium is made of two plates with narrow and acutely pointed anterior prongs. The cyrtomatodont teeth are short, thick and blunt. They are covered with secondary fibers. The hinge is straight. The smooth dorsal valve is lid-like with many concentric growth lines and has a posterior part slightly convex at the level of the circular protegulum that has a granulate surface. The peripheral rim is quite wide and clearly defined. The peripheral ridge is heavily tuberculate (Pl. 7, Fig. 1d). The median septum is thin, widening in its anterior part. The ventral surface of the septum is narrowly canaliculated and its lateral sides are irregularly spinous (Pl. 7, Fig. 1a, 1e). Brachial cavities (or brood pouches) are covered with canopies similar to spicular lattices leaving free interstices of irregular shape (Pl. 7, Fig. 1a–e). Canopying spicules are not very numerous and relatively thick with a finely granulated surface (Pl. 7, Fig. 1b). The cardinal process is curved backwards. Inner socket ridges are very thick and covered with secondary shell material. The brachial bridge, with a denticulate ventral crest, is convex posteriorly. Lophophore muscle scars are strong and clearly visible on its anterior surface (Pl. 7, Fig. 1d). Comparison with Minutella minuta (Cooper, 1981) and Minutella cf. minuta (in Simon & Hoffmann 2013) The specimens from Lembeh as all Pacific Minutella representatives (see Bitner 2009, p. 18, fig. 13A–I; Hoffmann and Lüter 2010, pp. 150–152) are extremely similar to the specimens of M. minima sensu stricto from Samper Bank, Madagascar, as described by Cooper (1981). Some details are however different between the “populations”. In Lembeh, spicules of the canopies are generally coarser. The calcitic pole has also a different outline (Pl. 7, Fig. 1e). It has very poorly developed lateral outgrowths and also a posterior outgrowth reduced to a tiny and sharp triangle (Pl. 7, Fig. 1f). The specimens of M. cf. minuta (Indonesia) from Donggala in the Strait of Makassar described in Simon & Hoffmann (2013) have a longer interarea with a narrower rugideltidium. They have a thin lamellar calcitic pole with a quite extended posterior outgrowth (Simon & Hoffmann 2013, pl. 1, fig. 4d, 4g–h). Their canopies are made with slender and more numerous spicules leaving more regular free cavities (pl. 1, fig. 4e; pl. 2, fig. 1a–b). In M. minuta (Cooper, 1981) from Samper Bank (see pp. 61–62) the interarea is generally shorter with a relatively wider rugideltidium (see Cooper 1981, pl. 6, figs. 27–28). The canopies of brood pouches have similar irregular cavities (see Cooper 1981, pl. 6, figs. 37–38, 40). Also, in M. minuta from Samper Bank (see Cooper 1981, pp. 61–62; pl. 6, figs. 27–40) the calcitic pole is thicker with a short posterior outgrowth but larger lateral outgrowths. However, these comparisons concern characters that remain affected by variability both in M. minuta (Cooper, 1981) and in all M. cf. minuta from Pacific origin. These variations should be studied statistically on very large samples in order to evaluate the value of such morphological features. But predominantly, living specimens of all “populations” of M. minuta and M. cf. minuta from the Indo-Pacific must be available to benefit from molecular investigations based on studies of the small subunit of the nuclear ribosomal RNA gene (18S rDNA). Ontogeny. The shell development for this species has been studied and illustrated in detail by Simon & Hoffmann (2013, p. 425–427; pl. 2, figs. 1–8) and can be consulted., Published as part of Simon, Eric, Lüter, Carsten, Logan, Alan & Mottequin, Bernard, 2018, Recent thecideide brachiopods (Thecideida, Thecideoidea) from northern Sulawesi (Indonesia) with discovery of a new Thecidellina species (Thecidellinidae), pp. 481-515 in Zootaxa 4526 (4) on pages 495-496, DOI: 10.11646/zootaxa.4526.4.4, http://zenodo.org/record/2611832, {"references":["Cooper, G. A. (1981) Brachiopoda from Southern Indian Ocean (Recent). Smithsonian Contributions to Paleobiology, 43, 1 - 93. https: // doi. org / 10.5479 / si. 00810266.43.1","Logan, A. & Baker, P. (2013) The development and shell microstructure of the pseudodeltidium and interarea in thecideide brachiopods. Palaeontology, 56, 433 - 455. https: // doi. org / 10.1111 / pala. 12001","Simon E. & Hoffmann, J. (2013) Discovery of Recent thecideide brachiopods (Order: Thecideida, Family: Thecideidae) in Sulawesi, Indonesian Archipelago, with implications for reproduction and shell size in the genus Ospreyella. Zootaxa, 3694 (5), 401 - 433. https: // doi. org / 10.11646 / zootaxa. 3694.5.1","Bitner, M. A. (2009) Recent Brachiopoda from the Norfolk Ridge, New Caledonia, with description of four new species. Zootaxa, 2235, 1 - 39.","Hoffmann, J. & Luter, C. (2010) Shell development in thecidellinine brachiopods with description of a new Recent genus. Special Papers in Palaeontology, 84, 137 - 160."]}

SUBPHYLUM RHYNCHONELLIFORMEA WILLIAMS ET AL., 1996 CLASS RHYNCHONELLATA WILLIAMS ET AL., 1996

Lenticellaria marerubris Simon, Logan & Mottequin sp. nov. Pl. 11, Figs. 1���6. 1921 Megerlia truncata var. paucistriata: Jackson, p. 48. 1973 Megerlia echinata (Fischer & Oehlert): Cooper, p. 18���19, pl. 3, figs. 27���31). 2008 Megerlia truncata var. paucistriata, Jackson: Bitner, Logan & Gischler, p. 279. pp 2008 Megerlia truncata: Bitner, Logan & Gischler, pp. 279, 284. pp v 2008 Megerlia echinata (Fischer & Oehlert, 1890): Logan, Toma��ov��ch, Zuschin & Grill, p. 395; fig. Q non figs. 2, P, R, S, T. pp v 2013 Megerlia echinata (Fischer & Oehlert): Logan & Bitner, pp. 161, 171. Holotype. Lenticellaria marerubris sp. nov. Pl. 11, Figs. 1 a��� 1 o. Well preserved articulated specimen that has been opened for the present study. It is preserved in the collections of the Natural History Museum of Vienna, Austria (NHMW 87192 /GP 247). The morphological measurements of the holotype are indicated in Table 2. Paratypes: NHMW 87192 /GP 248: small articulated adult (Pl. 11, Figs. 2 a���e; Table 2) with a dorso-biconvex structure. NHMW 87192 /GP 249: large articulated adult with a quite distorted outline (Pl. 11, Figs. 3 a���e; Table 2). NHMW 87192 /GP 250: medium sized adult specimen used for illustrating the ventral external ornamentation (Pl. 11, Fig. 4 a; Table 2). NHMW 87192 /GP 251: early juvenile articulated specimen with a loop at the early cucullate stage of development (Pl. 11, Figs. 5 a���f; Table 2). NHMW 87192 /GP 252: young articulated specimen showing the two claws of the loop at earliest stage of growth (Pl. 11, Figs. 6 a���d). Etymology. the specific name ��� marerubris ��� is the latin translation of ���from the Red Sea���. Locus typicus: Dahab, Egypt, Gulf of Aqaba at a depth of 12 m. Material investigated. Most of the specimens studied were collected by M. Zuschin (Fig. 4) in Dahab, Gulf of Aqaba, Egypt in very shallow water (12 m) and a few specimens more were collected at Port Sudan (see also Logan et al., 2008, fig. 1, p. 300 and Table 1, p. 301). For this study 38 complete articulated shells, 26 dorsal valves (sometimes fragmented) and 48 ventral valves (sometimes fragmented) were observed. Diagnosis of species. Micromorphic brachiopod with a very variable outline in dorsal view depending on the topography of the substrate to which the specimen is attached. Shell often distorted but sometimes semicircular in outline. External surface of dorsal valve smooth. Ventral valve with radial tuberculation. Between the radial rows very small scales are regularly developed in concentric lines. Foramen amphithyrid. Cardinal process and hinge plates absent. Inner dorsal valve floor tuberculate. Short crura developed. Loop made of two claws relatively narrow and thin with sharp tip and with a clearly raised pedestal. Descending branches absent. Septum on the ventral valve floor partly interrupted in its posterior portion. Diagnose de l���esp��ce. Brachiopode micro-morphique de forme tr��s variable en vue dorsale d��pendant de la structure du substrat sur laquelle le sp��cimen vit fix��. La coquille est souvent fortement d��form��e mais parfois elle peut appara��tre semi-circulaire et plus r��guli��re. La surface externe de la valve dorsale est lisse. La valve ventrale est radialement tubercul��e. Entre les rang��es de tubercules se d��veloppent de tr��s petites ��cailles dispos��es concentriquement. Foramen amphithyride. Processus cardinal et plaques de charni��res absentes. Surface interne de la valve dorsale tubercul��e. Crura tr��s courtes clairement d��velopp��es. Le brachidium se compose de deux pinces plut��t ��troites et minces avec des pointes aig��es. Le septum bifide est nettement sur��lev��. Branches descendantes absentes. Septum de la valve ventrale interrompu dans sa partie post��rieure. Mendiagnosia. �� Micromorphic �� Brachiopoda dengan bentuk yang sangat bervariasi dilihat atas katup dorsal.Kadang-kadang bentuk juga setengah lingkaran tergantung pada substrat dimana spesimen hidup dilampirkan. Shell sering sangat terdistorsi tapi kadang-kadang bisa muncul berbentuk setengah lingkaran dan lebih teratur. Permukaan luar katup dorsal halus sedangkan permukaan luar katup ventral adalah baris radial dengan tuberkel terucut yang kecil. Antara baris radial dari tuberkel ada banyak skala yang sangat kecil dikembangkan di garis konsentris. Amphithyrid foramen. Proses kardinal dan engsel piring tidak ada. Dalam lantai katup punggung ada tuberkel juga. Krura singkat dikembangkan. Brachidium merupakan dua cakar, sempit dan tipis sederhana dengan teratas tajam. Septum bercabang secara signifikan meningkat. Cabang turun tidak ada. Septum dari katup ventral terputus di bagian posteriornya. Description. Micromorphic kraussinid brachiopod (Table 2) with a very variable outline in dorsal view depending on the topography of the substrate to which the shell is attached. This species probably lives with its dorsal valve applied tightly to the substrate. The shell is plano-convex or concavo-convex, ventri-biconvex (pl. 11, Fig. 1 d), sometimes biconvex and rarely dorso-biconvex (Pl. 11, Fig. 2 c��� 2 d). Cardinal area orthocline. Anterior commissure rectimarginate, slightly sulcate to sulcate. Lateral commissure sinuous. Beak erect, often with intact obtuse umbo, sometimes slightly abraded. Amphithyrid foramen shared conspicuously by the dorsal valve. The dorsal side of the foramen is very convex (Pl. 11, Figs. 1g, 2 c, 2 e). Deltidial plates, very small, triangular, disjunct. Dorsal valve with external surface smooth except for several distinct growth lines. Ventral valve external surface with radial tuberculation and many very small scales disposed concentrically on the surface between the radial rows of tubercles (Pl. 11, Figs. 3 b, 4 a, 5 b, 6 b). The tubercles in this species are quite spiny in lateral view (Pl. 11, Figs. 1 d���e, 2 c). Obtuse hinge angle quite variable depending of the peculiar outline of each specimen. Cyrtomatodont teeth smooth, robust, short, flat dorsally and obtusely pointed. Pedicle collar thickened, sessile (Pl. 11, Fig. 1 n) with numerous punctae parallel to the external surface (visible with a binocular microscope, not with the SEM). Socket ridges thick, mainly the inner socket ridge that is also higher. The sockets are not deep. Cardinal process absent. Ventral valve with a low thick septum interrupted in its posterior part. The septum reaches the pedicle collar but does not join it. The commissural dorsal valve floor is heavily tuberculate. The commissural ventral valve floor is not tuberculate but hollow traces of the tips of dorsal tubercles are clearly visible. The brachidium consists of two pointed relatively slender claws attached to the dorsal valve floor by a relatively high pedestal. The distance between the tips of the claws is highly variable (Table 2) and depends partly on the general size of the specimen but also on the stage of ontogeny reached. Short crura are clearly developed but descending branches are never produced., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on pages 20-22, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667, {"references":["Fischer, P. & Oehlert, D. P. (1890) Diagnoses de nouveaux Brachiopodes. Journal de Conchyliologie, ser. 3, 38 (1), 70 - 74."]}

The genus Megerlia King, 1850 The main question of this paper is ���to be or not to be Megerlia ���. The type species of this genus is Megerlia truncata (Linn��, 1767). The original material described by Linn�� as Anomia truncata apparently consists of two dried specimens (as indicated in the Catalogue of Type specimens of the University of Uppsala: n�� 1907 a and n�� 1907 b) and should be Norwegian specimens (��� Habitat in Pelago Norvegico Supra corallia ��� Linn��, 1767; p. 1152, n�� 229). However, the presence of this species off Norway was not mentioned by Logan (2007) or Hiller et al. (2008). The species was not found in northerly latitudes such as the Irish coasts by Massy (1925), Norway by Wesenberg-Lund (1939), the North Sea (Cori, 1933) and the Faroe Islands (Thomsen, 2001). In the 19 th century Megerlia species were not recognized along the British coasts (Forbes & Hanley 1850, 1853). The original Linnaean specimens may have come from another area in the eastern Atlantic or even from the Mediterranean Sea. The current genus description provided by Lee & MacKinnon (2006, p. H 2245) stresses the large foramen which is often abraded (considered submesothyrid to amphithyrid), the disjunct deltidial plates and the tuberculate internal valve floors. No cardinal process is developed. Crural bases are present and attached to the inner sides of socket ridges. Descending branches join the cardinalia with the posterior part of the ring of the loop. The brachidium is typically bifurcate (see MacKinnon & Lee, 2006, fig. 1312) and a complete ring is built; this is unique among Kraussinoidea. SEM observations show in detail the development in M. truncata and interpretation of the earliest growth stages is here discussed again. The smallest specimen collected from Anzio has a width of 1.4 mm (Pl. 1, Figs. 1 a, 1 b). This specimen is slightly biconvex. The tip of the beak is abraded but the submesothyrid foramen is still intact on its dorsal side. The dorsal valve is smooth and the ventral valve presents some rare tubercles near the anterior commissure. The second specimen (Pl. 1, Figs. 2 a, 2 b) has a width of 1.73 mm. The shell becomes ventribiconvex with a smooth dorsal valve whereas the external surface of the ventral valve presents a radial series of tubercles. Noteworthy is that the outer shell surface between the radial ornamentation is smooth. The beak is more abraded and the dorsal valve is affected as the foramen seems amphithyrid. This gradual change is quite continuous during growth as seen on Pl. 1, Figs. 3���7. It appears that the foramen is more and more abraded due to the shortness of the pedicle and the high energy environment where M. truncata is generally living. Another development is the appearance of radial costae with tubercles on the external side of the dorsal valve. This tuberculation becomes more important during growth (Pl. 2, Figs. 1���3) although the ventral tubercles always remain more developed in size. An underestimated character in M. truncata is the absence, at all growth stages, of secondary tubercles, spines or scales in the ���intercostal��� zones. In these zones the shell remains smooth and only concentric growth lines are perceptible (Pls. 1���5). This is not the case in M. echinata (Pl. 10, Figs. 1, 2a���d) as strong spines cover the whole external surface of the ventral valve (see also Fischer & Oehlert, 1891, pl. 7, figs 13 a and b). Examination of many specimens of M. truncata from the whole Mediterranean Sea also clearly shows that both valves are provided with radial tuberculation, the ventral tuberculation always being stronger than the dorsal one. Specimens with abraded dorsal tubercles are found but an entirely smooth dorsal valve without any tuberculation has not been observed. This feature is important for distinguishing M. truncata from M. echinata (see below)., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on pages 7-8, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667, {"references":["King, W. (1850) A monograph of the Permian fossils of England. Palaeontographical Society Monograph, 3, xxxvii + 258 pp. http: // dx. doi. org / 10.5962 / bhl. title. 114608","Linne, C. A. (1767) Systema Naturae, sive Regna tria Naturae systematicae proposita per Classes, Ordines, Genera, Species cum Characteribus & Differentiis. 12 th edit., vol. 1, n ° 2, Holmiae, pp. 533 - 1327.","Hiller, N., MacKinnon, D. I. & Nielsen, S. N. (2008) A review of the systematic, biogeography and evolutionary relationships of Recent and fossil brachiopods of the Superfamily Kraussinoidea Dall, with descriptions of two new fossil species from New-Zealand and Chile. Earth and Environmental Science transactions of the Royal Society of Edinburgh, 98, 379 - 390. http: // dx. doi. org / 10.1017 / S 1755691007078474","Wesenberg-Lund, E. (1939) Recent Norwegian Brachiopods. Det Kongelige Norske Videnskabers-Selskab Forhandlinger, 11 (52), 201 - 204.","Cori, C. I. (1933) Brachiopoda. In: Grimpe, G & Wagler, E. (Eds.), Die Tierwelt der Nord-und Ostsee, 24 (VIIc 3): 132 - 149.","Thomsen, E. (2001) Brachiopoda in the Faroe Island area. Frodskaparrit, 49, 109 - 126."]}

Hillerella bisepta Simon, Logan & Mottequin, sp. nov. (Pl. 12. Figs. 1���7) Megerlia echinata (Fischer & Oehlert, 1890) ��� Logan, Toma��ov��ch, Zuschin & Grill, 2008, p. 305, Figs. 2 P, R, S, T (not Q). Diagnosis. Micromorphic kraussinoid, wider than long, biconvex, ventral valve with tuberculate radial costae; dorsal valve with less prominent radial ornamentation; ventral valve interior with single central septum not interrupted in its posterior part, hinge line with prominent teeth; dorsal valve hinge line with sockets but without cardinal process or crura; dorsal valve floor with usually a single commissural row of tubercles, dorsal valve interior with two prominent subparallel septa directed posteriorly from loop and slightly converging towards the hinge line, loop consisting of an arch supported by two slightly outwardly-inclined plates attached to valve floor, with a thin ventral bridge and with ventro-anterior expansions not strongly developed. Diagnose. Brachiopode kraussino��de micro-morphique, plus large que long, biconvexe; valve ventrale avec c��tes radiales tubercul��es; valve dorsale avec c��tes moins pro��minentes; valve ventrale avec septum central non interrompu dans sa partie post��rieure; ligne de charni��re avec des dents pro��minentes; ligne de charni��re de la valve dorsale avec sockets mais sans processus cardinal ni crura; surface interne de la valve dorsale avec deux septa subparall��les pro��minents dirig��s post��rieurement du brachidium jusqu����� la charni��re; brachidium consistant en une arche support��e par deux plaques divergentes attach��es �� la surface interne de la valve avec une partie ventrale extr��mement fine et avec des expansions ventro-ant��rieures peu d��velopp��es. Etymology. Specific epithet describes the diagnostic subparallel septa in the interior of the dorsal valve. Type locality (locus typicus). Station 100 / 8 at 90m depth off Port Sudan, Red Sea (18 �� 57.5 ��� N, 38 �� 15.8 ��� E) (Grill & Zuschin 2001; Logan et al., 2008, Fig. 1). Holotype. Ventral and dorsal valves of an articulated specimen, Pl. 12, Figs. 1 a���f from type locality, with dimensions indicated in Table 3 (NHMW 87192 /GP 253). Paratypes. (Pl. 12, Figs. 2���6) all from the type locality except Pl. 12, Fig. 7 which has been collected from Toliara (SW. of Madagascar). Measurements are indicated in Table 3. (NHMW 87192 /GP 254���255 and NBM��� 0 10321, NBM���008251). Material. Seven articulated specimens, 90 dorsal valves often fragmented and rarely intact, 88 ventral valves often fragmented too, all from the type locality 100 / 8 at 90 m depth off Port Sudan, Red Sea (Fig. 3 and see Grill & Zuschin 2001 and Logan et al., 2008 for details of exact location, substrate type and depth). Also a single bivalved specimen collected by Jacques Picard c. 1965 from dredged sediments in the vicinity of shallow coral reefs (to which the specimen was presumably attached) at Toliara, south-western Madagascar. Description. Shell small, wider than long, maximum width about 3 mm at mid-line, usually biconvex, but occasionally verging on plano-convex; shell surface of ventral valve costate (Pl. 12, Figs. 1 a,c), with between 10��� 14 radiating ribs, shallow, furnished with strong tubercles, some with spinose tips (Pl. 12, Fig. 1 c,d). Anterior commissure rectimarginate, occasionally sulcate. Hinge line orthocline, with prominent abraded beak (Pl. 12, Fig. 2 a) with large amphithyrid foramen. Shell surface of dorsal valve with less prominent radial costae, and with fewer tubercles (Pl. 12, Fig. 1 b). Ventral valve interior with prominent pedicle collar, strongly folded laterally when seen in anterior view (Pl. 12, Fig. 2 a). Narrow striated interarea, very small triangular disjunct deltidial plates and prominent teeth (Pl. 12, Fig. 2 b). Median septum extending anteriorly to about half length of valve (Pl. 12, Fig. 2 a). No tubercles. Shell endopunctate. Dorsal valve interior without cardinal process, outer socket ridges very low, much higher inner socket ridges excavate below but without crura, dental sockets deep. Strong divergent ridges sustain the sockets in the bottom of the valve (Pl. 12, Figs. 4, 5a���b). Two prominent parallel septa extend posteriorly on valve floor from about the midpoint, lengthening with age but present in even the smallest specimens (Pl, 12, fig. 3). Anterior ends of septa attached to two slightly outwardly-inclined narrow plates anchored to, and emerging upwards from, the valve floor (Pl. 12, Fig. 5 c). An ontogenetic sequence shows the gradual development of the loop from a very early stage (Pl. 12, Figs. 1 e��� f) to later mature stages (Pl. 12, Figs. 4, 5a���d, 6, 7a���b). Plates support an arch or cross-bar which, with ontogeny, gradually builds inward from the top of either plate to eventually join, completing the cross-bar, which bridges the gap and acts as a support for the lophophore. The lateral terminations of the cross-bar may be bulbous (Pl. 12, Figs. 6, 7a���b). Anterior margin of dorsal valve with a single row of tubercles (Pl. 12, Figs. 1 e, 5 a, b, 7 a), with excavated tips (Pl. 12, Fig. 1 e, 5 a, 7 a). Shell endopunctate (Pl. 12, Fig. 5 c). PLATE 12. Hillerella bisepta sp. nov. all collected from Port Sudan at 90 m. Station 100, Fig. 3, except specimen figured in fig. 7, which comes from Toleara, Madagascar. Fig. 1. Hillerella bisepta sp. nov. Holotype (NHMW 87192 /GP 253). 1 a: External view of ventral valve showing radial ornament of costae, tubercles and small spines. 1 b: External view of dorsal valve showing radial ornament of faint ribbing, no tubercles or spines. 1 c���d: Profile view of both valves attached, showing differences in ornament between the two valves. 1 e���f. Interior of dorsal valve. 1 e: early stage development of septal ridges and arch of loop of brachial skeleton. Note single row of marginal tubercles with excavated tips. 1 f: enlargement of brachial skeleton and septal ridges shown in previous figure. Fig. 2. Paratype 1. (NHMW 87192 /GP 254). 2 a; Interior of ventral valve, with pedicle collar, hinge area, teeth, median septum and tuberculate margin. 2 b: enlargement of part of hinge area (H) of previous specimen, showing tooth (T) and disjunct deltidial plate (D) outlined in black. Fig 3. Paratype 2. (NHMW 103331 - 5). Interior of early juvenile stage in ontogeny of dorsal valve, showing biseptal development preceding loop. Fig. 4. Paratype NBM- 010321. Interior of juvenile dorsal valve, anterior view of loop of immature form showing progressive development of arch of loop towards fusion. Fig. 5. Paratype 4. (NHMW 87192 /GP 255). 5 a: ventral view. 5 b: anterior view. 5 c: oblique lateral view. 5 d: detail of the ventral bridge. Note marginal tubercles with excavated tips in 5 a. Fig. 6. Paratype NHMW 103335. Detailed view of a mature loop with bulbous extremities. Hillerella bisepta sp. nov. Specimen collected from Toliara (SW Madagascar), c 1965, dredged by J. Picard (no depth data). Fig. 7. Paratype NBM-008251. Interior of a dorsal valve. 7 a: general ventral view showing brachidium and marginal tubercles with excavated tips. 7 b: thickened mature loop and bulbous extremities. Remarks. This species was briefly described and illustrated by Logan et al. (2008) from specimens collected by Zuschin and others from a wide variety of localities in the northern and central areas of the Red Sea, including the Port Sudan area. It was misidentified as Megerlia echinata (Fischer & Oehlert 1890) at that time. Consequently we have now established the new genus Hillerella. While this genus has some of the attributes of the other members of the family Kraussinidae (Kraussina, Megerlia, Megerlina and Pumilus (Lee & MacKinnon, 2006) it differs from other members of this family mainly in its small size typical of Red Sea brachiopods in general (Zuschin & Mayrhofer, 2009), the possession of subparallel septal ridges on the floor of the dorsal valve and the unique structure of the loop. Unlike Hillerella, Lenticellaria has a single central pillar with a bifurcate loop on either side to support the lophophore, while the separate claws never conjoin. The single example from Toliara, Madagascar (Pl. 12, Fig. 7 a, b) shows a thicker crossbar than typical Red Sea specimens, with more bulbous lateral terminations and the space between the underlying supports is narrower at the base, but nevertheless has the diagnostic subparallel septal ridges and loop structure to justify being assigned to H. bisepta. Homeomorphy and comparison with other kraussinid brachiopods. Homeomorphy is common among brachiopods and is often an important cause of misinterpretation. The most striking homeomorphy for Lenticellaria gregoryi sp. nov. is observed with the recently erected Annuloplatidia curiosa Bitner, 2014 (see Bitner, 2014, Fig. 9 A���H, p. 257). This micromorphic species is easily confused with a young Megerlia or with our new species. The brachidium is however typical of the genus Annuloplatidia Zezina, 1981. The homeomorphy between L. gregoryi sp. nov. and young different species of Megerlia (Pls. 1���5) is clearly visible. Controversies concerning the validity of certain Megerlia species are still discussed in the literature and most are due to homeomorphy. M. echinata (Fischer & Oehlert, 1890), for instance, is considered as an ecological variant of M. truncata by Logan (1979) and Logan et al. (2004). For Miocene specimens Bitner (1990, 1993) has the same opinion. For Emig et al. (2015) this species is a synonym of M. truncata. On the contrary M. echinata was regarded as a separate species by Brunton & Curry (1979), Cooper (1973 b, 1981 a, 1981 b), Brunton (1989) and Anad��n (1994). It is reconsidered again as a valid species by Logan et al. (2007, 2008). Detailed morphological observations of the type specimen and SEM illustrations have not been published until now (Plate 10). The type specimen shows all the characters proposed independently by Atkins (1961 a, p. 89) for her M. echinata material collected by the R.V. Sarsia (1958 & 1959) at depths between 556��� 882 m. The flat dorsal valve is smooth (without any radial costae) with numerous concentric growth lines (Pl. 10, Fig. 1) whereas the ventral valve is ornamented by long spines (Pl. 10, Figs. 2 a���b). All these spines are not only placed on radial costae but they can be developed on the whole surface of valve (Pl. 10, Fig. 2 c). Broken spines show a typical alveolar structure (Pl. 10, Fig. 2 d). The differences between the lophophore of M. echinata and M. truncata were discussed by Atkins (1961 b). A problem of zoogeographical order remains. The original specimen of M. echinata collected by ���Le Travailleur��� has been trawled off Cape Bojador (Sudan) between 640���782 m deep. The material studied by Atkins came from the western Atlantic. Phylogenetic analyses should establish definitively the taxonomic status of M. echinata and its relationship with other megerlinid species and also will confirm if the material from Cape Bojador and from the western Atlantic are the same species. Similar problems in regard to Megerlia gigantea (Deshayes, 1863) were discussed by Cooper (1981) and still no definitive solution has been found. For Bitner (2007, 2008) and Emig et al. (2015) M. gigantea is regarded as a synonym of M. truncata whereas it is a distinct species for Laurin (1997), Cooper (1981) and Gaspard (2003). The specimen illustrated by Laurin as M. echinata (1997, fig. 38, p. 452 and fig. 39, p. 453) is quite unusual. No complete description of this material is given and the illustrations consist of two drawings. His fig. 38 may be ���synthetic��� as no details of the tuberculation and of the ventral ornamentation are provided. The loop represented in fig. 39 is quite different from the loop seen in Megerlia at the same stage of growth (Pl. 4, Figs. 3 a���e and Pl. 5, Figs. 1 a���g). These drawings probably represent another genus. Recently, Bitner (2014, p. 258) placed these specimens in synonymy with her newly-erected species Annuloplatida curiosa from French Polynesia. She suggests that Laurin made a wrong interpretation of his material. However, this material should be re-studied again in regard to its taxonomic position. The specimen reported by D���Hondt (1987) is more problematic and this material should be revised because his determination is based only on Cooper���s description (1981) and no description and illustration of this single specimen was provided. Note that among the morphological characters drawn on the original figure of Deshayes (1863, p. 37, pl. 2, figs. 10, 11) no ���ring��� is visible. Only a bifurcate septum producing a fork is apparent which is joined by descending branches. The ring is probably broken. The specimen of Megerlia echinata illustrated by Cooper (1973, pl. 1, figs. 1���3) collected from southwest Cape of Good Hope has a width of 10 mm and is much larger than L. gregoryi sp. nov. The dorsal valve is smooth and the ventral valve has radial tuberculation. The surface between the radial rows of tubercles is covered with straight small spines which differ from the scales observed in L. gregoryi sp. nov. Megerlia representatives and also M. truncata appear to have an almost global distribution (Logan, 2007; Bitner, 2007, 2008). This species is common in the Mediterranean Sea and seems to be widely distributed in the Caribbean Sea, along the West African coast (though not as far south as Cape Verde as indicated in Logan, 1988), Canary Islands and Madeira (Logan, 1983 & Logan, 1988) and as far as Japan, Philippines, Hawaii and other Pacific zones (see references in Logan, 2007, p. H 3114). Such a wide distribution, interrupted by vast gaps, is difficult to explain, and is in need of revision. Specimens from the Persian Gulf were assigned to this species by Jackson (1921) under the name Megerlia truncata var. paucistriata (see also Bitner, 2008, p. 279 and Bitner et al., 2008). They are not illustrated but the description fits perfectly with L. marerubris sp. nov. We put M. truncata var. paucistriata in synonymy with L. marerubris sp. nov. Some specimens of M. truncata should be revisited, for instance the specimen from French Polynesia described by Bitner (2007, fig. 3, I���J). This is a small specimen but its loop is unknown as dried soft parts are present. Bleaching this material should be useful for knowing the brachidium structure to aid in the generic identification. A similar question concerns the specimen illustrated in Bitner (2008, fig. 19, D���E) as it is very small. Its attribution to Megerlia or Annuloplatidia should be reconsidered. Recently, Bitner (2014) suggested that all her described Megerlia specimens from Pacific waters are in fact Annuloplatidia curiosa Bitner, 2014 and she placed these Megerlia specimens in synonymy with this platidiid brachiopod. However, as Lenticellaria gregoriae sp. nov is also quite similar to Annuloplatidia externally, bleaching this material is necessary for knowing the real brachidium structure and to aid in the generic identification. Megerlia acrura Hiller, 1986 may be another problematic species from the eastern coast of South Africa. The external outline is similar to Megerlia species but the brachidium which develops a complete ring does not possess crura and descending branches. Studying the brachiopods collected from the BENTHEDI��� cruise in the Mozambique, Zezina (1987) suggested the name Megerlina gigantea (Deshayes, 1863) for a kraussinid specimen devoid of crura. She did not give any comments concerning the presence of a ring (or the presence of a broken ring?). Her description is quite short and no illustration is available for this material. Bitner (2007) suggested that this kraussinid brachiopod could be an M. acrura. This opinion has been adopted again by Hiller et al. (2008). To know if M. acrura should be placed in a new genus was considered "a moot point" by Hiller et al. (2008, p. 381)., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on pages 26-30, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667, {"references":["Fischer, P. & Oehlert, D. P. (1890) Diagnoses de nouveaux Brachiopodes. Journal de Conchyliologie, ser. 3, 38 (1), 70 - 74.","Logan, A., Tomasovych, A., Zuschin, M. & Grill, B. (2008) Recent brachiopods from the Red Sea and Gulf of Aden. Fossils and Strata, 54, 299 - 309. http: // dx. doi. org / 10.1017 / S 1755691007078449","Grill, B. & Zuschin, M. (2001) Modern shallow-to deep-water bivalve death assemblages in the Red Sea-ecology and biogeography. Palaeogeography, Palaeoclimatology, Palaeoecology, 168, 75 - 96. http: // dx. doi. org / 10.1016 / S 0031 - 0182 (00) 00250 - 9","Zuschin, M. & Mayrhofer, S. (2009) Brachiopods from cryptic coral reef habitats in the Northern Red Sea. Facies 55, 335 - 344. http: // dx. doi. org / 10.1007 / s 10347 - 009 - 0189 - 1","Bitner, M. A. (2014) Living Brachiopods from French Polynesia, Central Pacific, with Descriptions of Two New Species. Pacific Science, 68 (2), 245 - 265. http: // dx. doi. org / 10.2984 / 68.2.6","Zezina, O. N. (1981) Recent deep-sea Brachiopoda from the western Pacific. Galathea Report, 15, 7 - 20.","Logan, A. (1979) The Recent Brachiopoda of the Mediterranean Sea. Bulletin de l'Institut oceanographique de Monaco, 72, n ° 1434, 1 - 112.","Logan, A., Bianchi, C. L., Morri, C. & Zibrowius, H. (2004) The present-day Mediterranean brachiopod fauna: diversity, life habits, biogeography and paleobiogeography. Scientia Marina, 68 (Supplement 1), 163 - 170. http: // dx. doi. org / 10.3989 / scimar. 2004.68 s 1163","Bitner, M. A. (1990) Middle Miocene (Badenian) brachiopods from the Roztocse Hills, south-eastern Poland. Acta Geologica Polonica, 40, 129 - 157.","Bitner, M. A. (1993) Middle Miocene (Badenian) brachiopods from coral reefs of north-western Bulgaria. Acta Geologica Polonica, 43, 147 - 155.","Emig, C. (2015) Megerlia. In: Emig, C. C., Alvarez, F. & Bitner, M. A. (Eds.), World Brachiopoda database. World Register of Marine Species. Avaliable from: http: // marinespecies. org / aphia. php? p = taxdetails & id = 104045 (Accessed 21 Jun. 2015)","Brunton, C. H. C. & Curry, G. B. (1979) Synopses of the British Fauna, n ° 17. British Brachiopods. Published for the Linnean Society of London and The Brackish-Water Sciences Association by Academic Press (Ed.), London, New York, Toronto, Sydney and San Francisco, 64 pp.","Cooper, G. A. (1973) New Brachiopoda from the Indian Ocean. Smithsonian Contributions to Paleobiology, 6, 1 - 43. http: // dx. doi. org / 10.5479 / si. 00810266.16.1","Cooper, G. A. (1981 a) Brachiopoda from the Gulf of Gascogne, France (Recent). Smithsonian Contributions to Paleobiology, 44, 1 - 28. http: // dx. doi. org / 10.5479 / si. 00810266.44.1","Cooper, G. A. (1981 b) Brachiopoda from Southern Indian Ocean (recent). Smithsonian Contributions to Paleobiology, 43, 1 - 93. http: // dx. doi. org / 10.5479 / si. 00810266.43.1","Brunton, C. H. C. (1989) Some Brachiopods from the eastern Mediterranean Sea. Israel Journal of Zoology, 35, 151 - 169. [printed 1988]","Anadon, N. (1994) Braquiopodos actuales de la plataforma y talud continental de la costa central de Asturias (Norte de Espana). Boletin de la Real Sociedad Espanola de Historia Natural, (Biol.), 91 (1 - 4), 65 - 77.","Atkins, D. (1961 a) The generic position of the brachiopod Megerlia echinata (Fischer & Oehlert). Journal of the Marine Biological Association of the U. K., 41 (1), 89 - 94. http: // dx. doi. org / 10.1017 / S 0025315400001545","Atkins, D. (1961 b) The growth stages and adult structure of the lophophore of the brachiopods Megerlia truncata (L.) and M. echinata (Fischer & Oehlert). Journal of the Marine Biological Association of the U. K., 41 (1), 95 - 111. http: // dx. doi. org / 10.1017 / S 0025315400001557","Deshayes, G. P. (1863) Catalogue des Mollusques de l'ile de la Reunion (Bourbon). In: Maillard, L. (Ed.), Notes sur l'Ile de la Reunion (Bourbon), 2, Dentu (Ed.), Paris, 144 pp.","Bitner, M. A, (2007) Recent brachiopods from the Austral Islands, French Polynesia, South-Central Pacific. Zoosystema, 29 (3), 419 - 461.","Laurin, B. (1997) Brachiopodes recoltes dans les eaux de la Nouvelle-Caledonie et des iles Loyaute, Matthew et Chesterfield. In: Crosnier, A. (Ed.), Resultats des Campagnes MUSORSTOM, volume 18. Memoires du Museum national d'Histoire Naturelle, 176, 411 - 471.","Gaspard, D. (2003) Recent brachiopods collected during the \" SEAMOUNT 1 \" CRUISE off Portugal and the Ibero - Moroccan Gulf (Northeastern Atlantic) in 1987. Geobios, 36, 285 - 304. http: // dx. doi. org / 10.1016 / S 0016 - 6995 (03) 00033 - 0","Logan, A. (1988) Brachiopoda collected by CANCAP IV-VI Expeditions to the south-east North Atlantic. 1980 - 1982. Zoologische Mededelingen, 62 (5), 59 - 74.","Logan, A. (1983) Brachiopoda collected by CANCAP I-III Expeditions to the south-east North Atlantic. 1976 - 1978. Zoologische Mededelingen, 57 (18), 165 - 189.","Jackson, J. W. (1921) On the occurrence of Lusitanian brachiopods in the Persian Gulf. Annals and Magazine of Natural History, (9), 7, 40 - 49. http: // dx. doi. org / 10.1080 / 00222932108632488","Hiller, N., MacKinnon, D. I. & Nielsen, S. N. (2008) A review of the systematic, biogeography and evolutionary relationships of Recent and fossil brachiopods of the Superfamily Kraussinoidea Dall, with descriptions of two new fossil species from New-Zealand and Chile. Earth and Environmental Science transactions of the Royal Society of Edinburgh, 98, 379 - 390. http: // dx. doi. org / 10.1017 / S 1755691007078474"]}

Lenticellaria gregoryi Simon, Logan & Mottequin, sp. nov. Pl. 6, Figs 1���4; Pl. 7, Figs. 1���3; Pl. 8, Figs. 1���2; Pl. 9, Figs. 1���2. Holotype. Lenticellaria gregoryi Simon et al., Pl. 6, Figs. 3 a���c; Pl. 8, Figs. 1 a���g, 2 a���f; Pl. 9, Figs. 1 a���e. It is a perfectly preserved articulated specimen which has been opened for the present study and is preserved in the collections of the Royal Belgian Institute of Natural Sciences, RBINS-BT. 4. The morphological measurements of the holotype are indicated in Table 1. Paratypes. RBINS-BT. 5. A complete articulated specimen illustrated from Kodingareng, Makassar, Indonesia Pl. 6, Figs. 1 a���b, 4 a���g and Pl. 7, Fig. 1 a���f. RBINS-BT. 6. A ventral valve from Kodingareng, Makassar, Indonesia illustrated Pl. 6, Fig. 2; Pl. 7, Fig. 2 a���f. RBINS-BT. 8. A ventral valve from Tulamben, Bali, Indonesia illustrated Pl. 7, Fig. 3 a���c. RBINS ���BT. 7. Complete articulated juvenile specimen from Kodingareng, Makassar, Indonesia illustrated Pl. 9, Figs. 2 a���g. Etymology. This species is dedicated to the late Gregory Willems and his first name is used for the species name. Locus typicus: Tanker-shipwreck near Kodingareng Island (Spermonde Archipelago), Makassar, South Sulawesi, Indonesia (5 ��06��� 17.43 ��� S 119 �� 17 ���03.92���E). Material investigated From Makassar (Kodingareng wreck): 15 specimens. 10 complete articulate specimens of different sizes (one was a living specimen, now preserved in ethanol), two isolated dorsal valves with brachidium and three isolated ventral valves. From Tulamben (Bali): one articulated specimen, one ventral valve and three incomplete ventral valves. Diagnosis of species. Micromorphic brachiopod, wider than long with a semicircular outline in dorsal view. The external surface of the dorsal valve is smooth whereas the external surface of the ventral valve has radial tuberculation. Between the radial rows of tubercles, numerous scales with typical ���Gaussian curve��� outline are developed in concentric lines. Foramen amphithyrid. Cardinal process and hinge plates absent. Inner dorsal valve floor tuberculate. Short crura developed. Loop, going through a cucullate phase, a ���pseudo��� annular phase and finally consisting of two simple claws, relatively wide, with sharp tips. Base of the bifid septum unraised. Descending branches absent. Septum on the ventral valve floor partly interrupted in its posterior portion. Diagnose de l���esp��ce. Brachiopode micro-morphique, plus large que long avec un aspect semi-circulaire en vue dorsale. La surface ext��rieure de la valve dorsale est lisse tandis que la surface de la ventrale montre une ornementation radialement tubercul��e. Entre les rang��es de tubercules se d��veloppent des ��cailles distribu��es en lignes concentriques. Foramen amphithyride. Processus cardinal et plaques de charni��re absentes. Surface dorsale interne tubercul��e. Crura tr��s courtes toujours pr��sentes. Brachidium adulte consistant en deux pinces �� extr��mit��s pointues form��es par une phase cuccul��e ��voluant par une �� pseudo �� phase annulaire. Branches descendantes absentes. Septum de la valve ventrale interrompu dans sa partie post��rieure. Mendiagnosia. �� Micromorphic �� Brachiopoda, lebih lebar dari panjang dengan bentuk setengah lingkaran kelau dilihat atas katup dorsal. Permukaan luar katup dorsal halus sedangkan permukaan luar katup ventral adalah baris radial dengan duri tebal. Antara baris radial dari tuberkel ada banyak skala yang berbentuk �� kurva Gaussian �� dikembangkan di garis konsentris. Foramen amphithyrid. Tidak ada kardinal proses. Tidak ada engsel piring. Dalam lantai katup punggung ada tuberkel juga. Krura singkat dikembangkan. Brachidium, akan melalui fase cucullate, sebuah "pseudo" berbentuk gelang fase dan akhirnya merupakan dua cakar, relativ luas, sederhana dengan teratas tajam. Septum mendua, masih sangat datar. Cabang turun tak pernah dikembangkan. Septum di lantai katup ventral sebagian terputus di bagian posteriornya. Description. Micromorphic kraussinid brachiopod more or less semi-circular in dorsal outline, slightly wider than long, ventribiconvex (Pl. 6. Fig. 4 c) to planoconvex or sometimes concavoconvex (Pl. 9, Fig. 2 a; Table 1). Cardinal area orthocline. Anterior commissure rectimarginate in juveniles to very slightly sulcate in adults. Beak PLATE 6. Lenticellaria gregoryi sp. nov. Material collected by G. Willems from a tanker shipwreck at Kodingareng (Spermonde Archipelago, west of Makassar, South Sulawesi, Indonesia) at a depth of 30 m. Size of the specimens indicated with scale bars. Fig. 1. RBINS-BT. 5. Paratype. Articulated specimen photographed with a macro-objective on a Pentax K 20 D. The general aspect of the shell (1 a; 1 b) is homeomorphic with juvenile specimens of Megerlia or Annuloplatidia curiosa Bitner, 2015. The shell surface ornamentation is unclear as it is difficult to see if the shell surface is covered with spines or scales. Under a binocular microscope there appear to be numerous punctae (depending on the type of light). Fig. 2. RBINS-BT. 6. Paratype, a ventral valve photographed with a macro-objective on a Pentax K 20 D. The typical parallel disposition of the punctae in the sessile pedicle collar is clearly visible. Fig. 3. RBINS-BT. 4. Holotype photographed with a macro-objective on a Pentax K 20 D. 3 a: external ventral view of the ventral valve. 3 b: Internal aspect of the ventral valve with the sessile pedicle collar showing parallel disposition of the punctae. 3 c: Internal dorsal view showing the brachidium and the very small crura. Fig. 4. RBINS-BT. 5. Paratype. Articulated specimen observed with SEM at low magnification. External aspect of the shell is homeomorph with Megerlia. Thee shell is slightly dorsi-biconvex (4 c; 4 d; 4 e). The dorsal valve is smooth (4 a); The anterior commissure is slightly sulcate. The foramen is not abraded and is amphithyrid (4 a; 4g). The extremely reduced brachidium is made of two claws and of two small crura (4 f). PLATE 7. Lenticellaria gregoryi sp. nov. Material collected by G. Willems. Figs. 1 a���f and 2 a���f: specimens from a shipwreck at Kodingareng (Spermonde Archipelago, west of Makassar, South Sulawesi, Indonesia) at a depth of 30 m. Figs. 3 a���c: specimen from the ���Liberty��� shipwreck at Tulamben (Bali Sea, Bali, Indonesia) at a depth of 25 m. Size of the specimens indicated with scale bars. Fig. 1. RBINS-BT. 5. Paratype. Articulated specimen observed with SEM at high magnification. External aspect of the shell. Radial rows of thick tubercles with shell surface between these rows of tubercles covered with small scales giving a ���lenticel��� aspect. At still higher magnification (1 b��� 1 f) the scales seen in anterior view appear as ���Gaussian curves��� (1 e). a: dorsal view; b: anterior view; c: posterior view; d: dorsal view at very high magnification; e: anterior view at very high magnification; f: posterior view at very high magnification. Fig. 2. RBINS-BT. 6. Paratype. Ventral valve. 2 a: general ventral view; 2 b: general internal view with sessile pedicle collar and low median septum interrupted in its posterior part; 2 c: detail of external ornamentation with numerous small scales; 2 d: detailed view of the sessile pedicle collar (with SEM the punctae are not visible); 2 e; detailed view of the posterior part of the septum. Fig. 3. RBINS-BT. 8. Paratype. Ventral valve. 3 a: general ventral view; 3 b: Scales ornamentation of shell surface at high magnification; 3 c: general internal view with sessile pedicle collar and low septum interrupted in its posterior part. PLATE 8. RBINS-BT. 4. Lenticellaria gregoryi sp. nov. Holotype collected by G. Willems from sieved sediment from a tanker shipwreck at Kodingareng (Spermonde Archipelago, west of Makassar, South Sulawesi, Indonesia) at a depth of 30 m. Detailed SEM views of both separated dorsal (Figs. 1 a���g) and ventral (Figs. 2 a���f) valves. Size of the specimen indicated with scale bars. Fig. 1. 1 a: Internal view of the dorsal valve showing subcircular tubercles on the valve floor, short intact crura and symmetrical sharply pointed claws; b���c: detailed view of the brachidium in ventral view. The sharp tips of the loop are clearly visible as the crura. Cardinal process absent (1 b). 1 c: detailed view of the loop at high magnification in ventral view. 1 d: Oblique lateral view; 1 e Detailed view of the loop in anterior view 1 f: Detailed view of the loop in lateral view; 1g detailed posterior view of the loop. (1 e; 1g) show also that resorption process of cucullate structure has begun anteriorly; 1 h: detailed view of the right crus; 1 i: detailed view of the left crus. Fig. 2. 2 a: general view of the ventral valve in ventral view. Homeomorphy with Megerlia species is evident. 2 b: anterior general view showing a ���granulate��� shell surface. 2 c���f: Detailed SEM views of the external ornamentation of the shell surface. Numerous scales with Gaussian curve aspect visible together with larger tubercles. 2 c���d: ventral view at moderate and higher magnification. 2 e: anterior view; 2 f: oblique left lateral view of left part of the shell. Low costae ornamented with tubercles are visible. PLATE 9. Lenticellaria gregoryi sp. nov. Material collected by G. Willems from sieved sediment from a tanker shipwreck at Kodingareng (Spermonde Archipelago, west of Makassar, South Sulawesi, Indonesia) at a depth of 30 m. Detailed SEM views of separated ventral valve of the holotype (Figs. 1 ���ae) and illustration of an early juvenile articulated paratype (Figs. 2 a���g).. Size of the specimen indicated with scale bars. Figs. 1. RBINS-BT. 4. Holotype. 1 a: general internal view of ventral valve in dorsal view. 1 b: detailed view sessile pedicle collar and posterior interrupted portion of the ventral septum. 1 c: detailed view of the cyrtomatodont tooth. 1 d: general oblique lateral view illustrating the low ventral septum. 1 e: anterior view of ventral valve. Figs. 2. RBINS-BT. 7. Paratype. 2 a: dorsal view of the articulated specimen. Dorsal valve smooth and foramen amphithyrid. 2 b: Ventral view with rows of tubercles. 2 c: detailed view of the ventral shell surface with scales between the radial rows of tubercles. 2 d: high SEM magnification of the ornamentation of ventral shell surface. 2 e: Opened articulated paratype showing the tuberculation of the dorsal valve floor and the brachidium. 2 f: detailed view of the brachidium in dorsal view. Resorption of the anterior portion of the cucullate phase is started. Crura are appearing as very small knots. 2g: detailed anterior view of the cucullate phase showing the importance of the anterior resorption process at this stage of growth. erect with obtuse umbo often abraded. Foramen amphithyrid, shared conspicuously by the dorsal valve in all stages of growth with convex dorsal side (Pl. 6, Figs. 4 e���g). Deltidial plates poorly developed when preserved, elongate triangular, very narrow, disjunct. Dorsal valve always smooth. On the contrary, the ventral valve shows a clear radial tuberculation from the juvenile stage of growth (Pl. 6, Fig. 4 b; pl. 7, Figs. 2 a, 3 a; Pl. 8, Figs. 2 a���c; Pl. 9, Fig. 2 b). A very different aspect of the ventral valve is observed with a SEM used at a greater magnification. The structure is reminiscent of wood (Pl. 7, Fig. 3 b). Spaces between the tuberculate radial rows are not smooth but ornamented with ���stripes��� oriented like the concentric growth lines. This is a striking distinction between this brachiopod and representatives of the genus Megerlia with a smooth or spiny shell surface between the tuberculate radial rows. However this morphological character is not easily seen with a binocular microscope and even with a SEM at low magnification (Pl. 7, Fig. 2 a, and Pl. 7, Fig. 3 a,). At still higher magnification, these ���stripes��� in dorsal view (Pl. 7, Figs. 1 a, 1 d, 2 c, 3 b) appear as minuscule scales with a ���Gaussian curve��� aspect in anterior view (Pl. 7, Fig. 1 b, 1 e, 1 f). Scales are densely present on all specimens found even on early juvenile shells (Pl. 9, Fig. 2 b). These scales with ���Gaussian curve��� outline are different from the tubercles which are more similar to conical relief. Obtuse hinge angle between 120 �� and 130 ��. Cyrtomatodont teeth curved, sharply ended and with a smooth surface. (Pl. 7, Fig. 2 d). Pedicle collar narrow and sessile (Pl. 7, Fig. 2 f) fused with the underlying shell and appearing as a narrow thickening of the shell (Pl. 7, Fig. 2 f). Shell endopunctate. Punctae of the pedicle collar parallel to the valve floor surface and perpendicular to the pores of the shell itself. This gives a characteristic striated aspect when observed with a binocular microscope (Pl. 6, Fig. 2). The number of punctae in the ventral valve varies around 294 punctae/mm ��. Ventral valve with a low thin median septum always interrupted in its posterior part (Pl. 7, Fig. 2 b, 2 f, 3 c; Pl. 9, Fig. 1 b, 1 e). This septum does not extend under the pedicle collar. Inner dorsal valve floor radially tuberculate. Inner ventral valve floor not tuberculate. Loop structure and ontogeny. The loop in the largest specimen is made of two crura and two claws only. This is always the case even in the most juvenile stage observed (Pl. 9, Figs. 2 e���g). During growth, these claws widens and the space between the tip of the claws increases (Pl. 6, Fig. 4 f; Pl. 8, Figs. 1 b, 1 e) but not regularly or linearly (Table 1). Crural bases are produced at an early stage of growth (Pl. 9, Fig. 2 f) but crura do not grow significantly: they remain short. Crural processes and descending branches are never produced. The ontogeny of this reduced brachidium begins as a typical Megerliinae process. The bifurcate septal pillar builds a posteriorly concave plate which becomes more conical. At this first stage the similarity with the ontogeny of Megerlia is evident. The resorption of the anterior part of this cucullate structure occurs at and just after the beginning. This resorption rapidly affects the anterior part of the cone and it continues posteriorly (Pl. 9, Fig. 2g). On the contrary, in Megerlia the resorption process begins posteriorly and stops before the anterior part could be dissolved (Pl. 3, Fig. 3 e), building a ring. Later in L. gregoryi sp. nov. when the resorption process is finished it remains two claws which increase in size progressively through growth but which never conjoin to form a ring (Table 1). This is one of the most reduced loop developments observed in Kraussinoidea. There are no ring and descending branches like in Megerlia. Crural bases are developed but ��� pro partim ��� descending branches extending from the septum like flanges, as seen in Megerlina, are not produced. The wide Y-shaped septum of Kraussina is quite different. The loop of Pumilus antiquatus Atkins, 1958 is the nearest more or less comparable structure. However, all other morphological features in P. antiquatus are markedly different (Lee and MacKinnon, 2006)., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on pages 14-20, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667, {"references":["Bitner, M. A. & Cohen, B. L. (2015) Congruence and conflict: case studies of morphotaxonomy versus rDNA gene tree phylogeny among articulate brachiopods (Brachiopoda: Rhynchonelliformea), with description of a new genus. Zoological Journal of the Linnean Society, 173, 486 - 504. http: // dx. doi. org / 10.1111 / zoj. 12217","Atkins, D. (1958) A new species and genus of Kraussinidae (Brachiopoda) with a note on feeding. Proceedings of the Zoological Society of London, 131 (4), 559 - 581. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1958. tb 00702. x"]}

Genus Lenticellaria Simon, Logan & Mottequin, gen. nov. Type species: Lenticellaria gregoryi Simon, Logan & Mottequin sp. nov. Etymology. from the word ���lenticel��� a botanical term meaning a kind of pore present on the surface of the bark of trees. Lenticels may be streak-like in shape. This name is derived from the scaly ornamentation of the shell surface. Diagnosis. Micromorphic kraussinoid slightly wider than long, with dorsal valve external surface smooth and radial tuberculate ventral valve. External surface of the ventral valve densely ornamented with very small scales disposed in concentric lines. Loop bifurcate, extremely reduced to two sharp claws. Crura present. Cardinal process, hinge plates and descending branches absent., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on page 14, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667

Hillerella bisepta Simon, Logan & Mottequin, sp. nov. (Pl. 12. Figs. 1–7) Megerlia echinata (Fischer & Oehlert, 1890) — Logan, Tomašových, Zuschin & Grill, 2008, p. 305, Figs. 2 P, R, S, T (not Q). Diagnosis. Micromorphic kraussinoid, wider than long, biconvex, ventral valve with tuberculate radial costae; dorsal valve with less prominent radial ornamentation; ventral valve interior with single central septum not interrupted in its posterior part, hinge line with prominent teeth; dorsal valve hinge line with sockets but without cardinal process or crura; dorsal valve floor with usually a single commissural row of tubercles, dorsal valve interior with two prominent subparallel septa directed posteriorly from loop and slightly converging towards the hinge line, loop consisting of an arch supported by two slightly outwardly-inclined plates attached to valve floor, with a thin ventral bridge and with ventro-anterior expansions not strongly developed. Diagnose. Brachiopode kraussinoïde micro-morphique, plus large que long, biconvexe; valve ventrale avec côtes radiales tuberculées; valve dorsale avec côtes moins proéminentes; valve ventrale avec septum central non interrompu dans sa partie postérieure; ligne de charnière avec des dents proéminentes; ligne de charnière de la valve dorsale avec sockets mais sans processus cardinal ni crura; surface interne de la valve dorsale avec deux septa subparallèles proéminents dirigés postérieurement du brachidium jusqu’à la charnière; brachidium consistant en une arche supportée par deux plaques divergentes attachées à la surface interne de la valve avec une partie ventrale extrêmement fine et avec des expansions ventro-antérieures peu développées. Etymology. Specific epithet describes the diagnostic subparallel septa in the interior of the dorsal valve. Type locality (locus typicus). Station 100 / 8 at 90m depth off Port Sudan, Red Sea (18 º 57.5 ’ N, 38 º 15.8 ’ E) (Grill & Zuschin 2001; Logan et al., 2008, Fig. 1). Holotype. Ventral and dorsal valves of an articulated specimen, Pl. 12, Figs. 1 a–f from type locality, with dimensions indicated in Table 3 (NHMW 87192 /GP 253). Paratypes. (Pl. 12, Figs. 2–6) all from the type locality except Pl. 12, Fig. 7 which has been collected from Toliara (SW. of Madagascar). Measurements are indicated in Table 3. (NHMW 87192 /GP 254–255 and NBM— 0 10321, NBM—008251). Material. Seven articulated specimens, 90 dorsal valves often fragmented and rarely intact, 88 ventral valves often fragmented too, all from the type locality 100 / 8 at 90 m depth off Port Sudan, Red Sea (Fig. 3 and see Grill & Zuschin 2001 and Logan et al., 2008 for details of exact location, substrate type and depth). Also a single bivalved specimen collected by Jacques Picard c. 1965 from dredged sediments in the vicinity of shallow coral reefs (to which the specimen was presumably attached) at Toliara, south-western Madagascar. Description. Shell small, wider than long, maximum width about 3 mm at mid-line, usually biconvex, but occasionally verging on plano-convex; shell surface of ventral valve costate (Pl. 12, Figs. 1 a,c), with between 10– 14 radiating ribs, shallow, furnished with strong tubercles, some with spinose tips (Pl. 12, Fig. 1 c,d). Anterior commissure rectimarginate, occasionally sulcate. Hinge line orthocline, with prominent abraded beak (Pl. 12, Fig. 2 a) with large amphithyrid foramen. Shell surface of dorsal valve with less prominent radial costae, and with fewer tubercles (Pl. 12, Fig. 1 b). Ventral valve interior with prominent pedicle collar, strongly folded laterally when seen in anterior view (Pl. 12, Fig. 2 a). Narrow striated interarea, very small triangular disjunct deltidial plates and prominent teeth (Pl. 12, Fig. 2 b). Median septum extending anteriorly to about half length of valve (Pl. 12, Fig. 2 a). No tubercles. Shell endopunctate. Dorsal valve interior without cardinal process, outer socket ridges very low, much higher inner socket ridges excavate below but without crura, dental sockets deep. Strong divergent ridges sustain the sockets in the bottom of the valve (Pl. 12, Figs. 4, 5a–b). Two prominent parallel septa extend posteriorly on valve floor from about the midpoint, lengthening with age but present in even the smallest specimens (Pl, 12, fig. 3). Anterior ends of septa attached to two slightly outwardly-inclined narrow plates anchored to, and emerging upwards from, the valve floor (Pl. 12, Fig. 5 c). An ontogenetic sequence shows the gradual development of the loop from a very early stage (Pl. 12, Figs. 1 e– f) to later mature stages (Pl. 12, Figs. 4, 5a–d, 6, 7a–b). Plates support an arch or cross-bar which, with ontogeny, gradually builds inward from the top of either plate to eventually join, completing the cross-bar, which bridges the gap and acts as a support for the lophophore. The lateral terminations of the cross-bar may be bulbous (Pl. 12, Figs. 6, 7a–b). Anterior margin of dorsal valve with a single row of tubercles (Pl. 12, Figs. 1 e, 5 a, b, 7 a), with excavated tips (Pl. 12, Fig. 1 e, 5 a, 7 a). Shell endopunctate (Pl. 12, Fig. 5 c). PLATE 12. Hillerella bisepta sp. nov. all collected from Port Sudan at 90 m. Station 100, Fig. 3, except specimen figured in fig. 7, which comes from Toleara, Madagascar. Fig. 1. Hillerella bisepta sp. nov. Holotype (NHMW 87192 /GP 253). 1 a: External view of ventral valve showing radial ornament of costae, tubercles and small spines. 1 b: External view of dorsal valve showing radial ornament of faint ribbing, no tubercles or spines. 1 c–d: Profile view of both valves attached, showing differences in ornament between the two valves. 1 e–f. Interior of dorsal valve. 1 e: early stage development of septal ridges and arch of loop of brachial skeleton. Note single row of marginal tubercles with excavated tips. 1 f: enlargement of brachial skeleton and septal ridges shown in previous figure. Fig. 2. Paratype 1. (NHMW 87192 /GP 254). 2 a; Interior of ventral valve, with pedicle collar, hinge area, teeth, median septum and tuberculate margin. 2 b: enlargement of part of hinge area (H) of previous specimen, showing tooth (T) and disjunct deltidial plate (D) outlined in black. Fig 3. Paratype 2. (NHMW 103331 - 5). Interior of early juvenile stage in ontogeny of dorsal valve, showing biseptal development preceding loop. Fig. 4. Paratype NBM- 010321. Interior of juvenile dorsal valve, anterior view of loop of immature form showing progressive development of arch of loop towards fusion. Fig. 5. Paratype 4. (NHMW 87192 /GP 255). 5 a: ventral view. 5 b: anterior view. 5 c: oblique lateral view. 5 d: detail of the ventral bridge. Note marginal tubercles with excavated tips in 5 a. Fig. 6. Paratype NHMW 103335. Detailed view of a mature loop with bulbous extremities. Hillerella bisepta sp. nov. Specimen collected from Toliara (SW Madagascar), c 1965, dredged by J. Picard (no depth data). Fig. 7. Paratype NBM-008251. Interior of a dorsal valve. 7 a: general ventral view showing brachidium and marginal tubercles with excavated tips. 7 b: thickened mature loop and bulbous extremities. Remarks. This species was briefly described and illustrated by Logan et al. (2008) from specimens collected by Zuschin and others from a wide variety of localities in the northern and central areas of the Red Sea, including the Port Sudan area. It was misidentified as Megerlia echinata (Fischer & Oehlert 1890) at that time. Consequently we have now established the new genus Hillerella. While this genus has some of the attributes of the other members of the family Kraussinidae (Kraussina, Megerlia, Megerlina and Pumilus (Lee & MacKinnon, 2006) it differs from other members of this family mainly in its small size typical of Red Sea brachiopods in general (Zuschin & Mayrhofer, 2009), the possession of subparallel septal ridges on the floor of the dorsal valve and the unique structure of the loop. Unlike Hillerella, Lenticellaria has a single central pillar with a bifurcate loop on either side to support the lophophore, while the separate claws never conjoin. The single example from Toliara, Madagascar (Pl. 12, Fig. 7 a, b) shows a thicker crossbar than typical Red Sea specimens, with more bulbous lateral terminations and the space between the underlying supports is narrower at the base, but nevertheless has the diagnostic subparallel septal ridges and loop structure to justify being assigned to H. bisepta. Homeomorphy and comparison with other kraussinid brachiopods. Homeomorphy is common among brachiopods and is often an important cause of misinterpretation. The most striking homeomorphy for Lenticellaria gregoryi sp. nov. is observed with the recently erected Annuloplatidia curiosa Bitner, 2014 (see Bitner, 2014, Fig. 9 A–H, p. 257). This micromorphic species is easily confused with a young Megerlia or with our new species. The brachidium is however typical of the genus Annuloplatidia Zezina, 1981. The homeomorphy between L. gregoryi sp. nov. and young different species of Megerlia (Pls. 1–5) is clearly visible. Controversies concerning the validity of certain Megerlia species are still discussed in the literature and most are due to homeomorphy. M. echinata (Fischer & Oehlert, 1890), for instance, is considered as an ecological variant of M. truncata by Logan (1979) and Logan et al. (2004). For Miocene specimens Bitner (1990, 1993) has the same opinion. For Emig et al. (2015) this species is a synonym of M. truncata. On the contrary M. echinata was regarded as a separate species by Brunton & Curry (1979), Cooper (1973 b, 1981 a, 1981 b), Brunton (1989) and Anadón (1994). It is reconsidered again as a valid species by Logan et al. (2007, 2008). Detailed morphological observations of the type specimen and SEM illustrations have not been published until now (Plate 10). The type specimen shows all the characters proposed independently by Atkins (1961 a, p. 89) for her M. echinata material collected by the R.V. Sarsia (1958 & 1959) at depths between 556– 882 m. The flat dorsal valve is smooth (without any radial costae) with numerous concentric growth lines (Pl. 10, Fig. 1) whereas the ventral valve is ornamented by long spines (Pl. 10, Figs. 2 a–b). All these spines are not only placed on radial costae but they can be developed on the whole surface of valve (Pl. 10, Fig. 2 c). Broken spines show a typical alveolar structure (Pl. 10, Fig. 2 d). The differences between the lophophore of M. echinata and M. truncata were discussed by Atkins (1961 b). A problem of zoogeographical order remains. The original specimen of M. echinata collected by “Le Travailleur” has been trawled off Cape Bojador (Sudan) between 640–782 m deep. The material studied by Atkins came from the western Atlantic. Phylogenetic analyses should establish definitively the taxonomic status of M. echinata and its relationship with other megerlinid species and also will confirm if the material from Cape Bojador and from the western Atlantic are the same species. Similar problems in regard to Megerlia gigantea (Deshayes, 1863) were discussed by Cooper (1981) and still no definitive solution has been found. For Bitner (2007, 2008) and Emig et al. (2015) M. gigantea is regarded as a synonym of M. truncata whereas it is a distinct species for Laurin (1997), Cooper (1981) and Gaspard (2003). The specimen illustrated by Laurin as M. echinata (1997, fig. 38, p. 452 and fig. 39, p. 453) is quite unusual. No complete description of this material is given and the illustrations consist of two drawings. His fig. 38 may be “synthetic” as no details of the tuberculation and of the ventral ornamentation are provided. The loop represented in fig. 39 is quite different from the loop seen in Megerlia at the same stage of growth (Pl. 4, Figs. 3 a–e and Pl. 5, Figs. 1 a–g). These drawings probably represent another genus. Recently, Bitner (2014, p. 258) placed these specimens in synonymy with her newly-erected species Annuloplatida curiosa from French Polynesia. She suggests that Laurin made a wrong interpretation of his material. However, this material should be re-studied again in regard to its taxonomic position. The specimen reported by D’Hondt (1987) is more problematic and this material should be revised because his determination is based only on Cooper’s description (1981) and no description and illustration of this single specimen was provided. Note that among the morphological characters drawn on the original figure of Deshayes (1863, p. 37, pl. 2, figs. 10, 11) no “ring” is visible. Only a bifurcate septum producing a fork is apparent which is joined by descending branches. The ring is probably broken. The specimen of Megerlia echinata illustrated by Cooper (1973, pl. 1, figs. 1–3) collected from southwest Cape of Good Hope has a width of 10 mm and is much larger than L. gregoryi sp. nov. The dorsal valve is smooth and the ventral valve has radial tuberculation. The surface between the radial rows of tubercles is covered with straight small spines which differ from the scales observed in L. gregoryi sp. nov. Megerlia representatives and also M. truncata appear to have an almost global distribution (Logan, 2007; Bitner, 2007, 2008). This species is common in the Mediterranean Sea and seems to be widely distributed in the Caribbean Sea, along the West African coast (though not as far south as Cape Verde as indicated in Logan, 1988), Canary Islands and Madeira (Logan, 1983 & Logan, 1988) and as far as Japan, Philippines, Hawaii and other Pacific zones (see references in Logan, 2007, p. H 3114). Such a wide distribution, interrupted by vast gaps, is difficult to explain, and is in need of revision. Specimens from the Persian Gulf were assigned to this species by Jackson (1921) under the name Megerlia truncata var. paucistriata (see also Bitner, 2008, p. 279 and Bitner et al., 2008). They are not illustrated but the description fits perfectly with L. marerubris sp. nov. We put M. truncata var. paucistriata in synonymy with L. marerubris sp. nov. Some specimens of M. truncata should be revisited, for instance the specimen from French Polynesia described by Bitner (2007, fig. 3, I–J). This is a small specimen but its loop is unknown as dried soft parts are present. Bleaching this material should be useful for knowing the brachidium structure to aid in the generic identification. A similar question concerns the specimen illustrated in Bitner (2008, fig. 19, D–E) as it is very small. Its attribution to Megerlia or Annuloplatidia should be reconsidered. Recently, Bitner (2014) suggested that all her described Megerlia specimens from Pacific waters are in fact Annuloplatidia curiosa Bitner, 2014 and she placed these Megerlia specimens in synonymy with this platidiid brachiopod. However, as Lenticellaria gregoriae sp. nov is also quite similar to Annuloplatidia externally, bleaching this material is necessary for knowing the real brachidium structure and to aid in the generic identification. Megerlia acrura Hiller, 1986 may be another problematic species from the eastern coast of South Africa. The external outline is similar to Megerlia species but the brachidium which develops a complete ring does not possess crura and descending branches. Studying the brachiopods collected from the BENTHEDI– cruise in the Mozambique, Zezina (1987) suggested the name Megerlina gigantea (Deshayes, 1863) for a kraussinid specimen devoid of crura. She did not give any comments concerning the presence of a ring (or the presence of a broken ring?). Her description is quite short and no illustration is available for this material. Bitner (2007) suggested that this kraussinid brachiopod could be an M. acrura. This opinion has been adopted again by Hiller et al. (2008). To know if M. acrura should be placed in a new genus was considered "a moot point" by Hiller et al. (2008, p. 381).

Genus Hillerella Simon, Logan & Mottequin gen. nov. Type species Hillerella bisepta Simon, Logan & Mottequin sp. nov. Etymology. Named for Norton Hiller in recognition of his important studies on Recent kraussinoid brachiopods. Diagnosis. Micromorphic kraussinoid, wider than long, biconvex, tuberculate radial costae mainly on ventral valve; dorsal valve with less prominent costae; ventral valve interior with single central septum, dorsal valve without cardinal process or crura; floor of dorsal valve interior with two prominent subparallel septa directed posteriorly from loop towards the hinge line; loop consisting of an arch supported by two slightly divergent plates attached to valve floor. Diagnose. Brachiopode kraussino��de micro-morphique, plus large que long, biconvexe; valve ventrale avec c��tes radiales tubercul��es; valve dorsale avec c��tes moins pro��minentes; valve ventrale avec septum central; valve dorsale sans processus cardinal ni crura; surface interne de la valve dorsale avec deux septa subparall��les pro��minents dirig��s post��rieurement du brachidium jusqu����� la charni��re; brachidium consistant en une arche support��e par deux plaques divergentes attach��es �� la surface interne de la valve. PLATE 11. Lenticellaria marerubris sp. nov. Material collected from the Red Sea, Gulf of Akaba, Dahab, by Martin Zuschin (NHMW) at a depth of 12 m. Size of the specimens indicated with scale bars. Fig. 1. Holotype. Articulated fully adult specimen opened for this study in order to illustrate the brachidium (NHMW 87192 / GP 247). 1 a: dorsal view showing the smooth dorsal valve with concentric growth lines and the wide amphithyrid foramen. This specimen is not distorted and has a nearly semicircular outline. 1 b: ventral view showing the tuberculated surface of the ventral valve. Radial rows of tubercles are clearly visible and between these rows an ornamentation made of many concentric lines of very small scales. 1 c: detailed view of the tubercles and scales of the same ventral valve. 1 d: lateral view. The beak is erect. The lateral commissure is sinuous. In this position the pointed tubercles and the scaly surface of the ventral valve are clearly visible. 1 e: detailed view of the lateral view. 1 f: anterior view. Anterior commissure is rectimarginate in this specimen. 1g: posterior view. The strong convexity of the dorsal side of the foramen is typical for all specimens investigated. 1 h: ventral view of the tuberculate dorsal valve showing the brachidium developing two claws supported by a pedestal. Short crura are visible. 1 i: internal anterior view of the dorsal valve showing the high pedestal, the two slender claws and the short crura. The crura are not supported by crural plates or thick crural ridges. 1 j: oblique lateral view. 1 k: oblique antero-lateral view. 1 l: detailed view of the short crura. Fig. 2. Paratype. Articulated fully adult specimen presented for illustrating the possibility of dorso-biconvex shell and distortion. (NHMW 87192 /GP 248). 2 a: dorsal view showing smooth dorsal valve and large amphithyrid foramen. Note the cordate outline for this specimen. 2 b: ventral view showing the radial rows of tubercles and the scaly surface of the valve. 2 c: lateral view showing the sinuous lateral commissure, the convex dorsal of the foramen and the spiny surface of the ventral valve. 2 d: anterior view. 2 e: posterior view. Fig. 3. Paratype. Articulated fully adult specimen used for illustrating a strongly distorted shell (NHMW 87192 /GP 249). 3 a: internal view of the tuberculate dorsal valve with brachidium. 3 b: detailed view of the external ventral valve ornamentation. 3 c: detail ventral view of the brachidium with its pedestal. 3 d: anterior view of the two claws of the brachidium. 3 e: detailed view of crura. Fig. 4. Paratype. Articulated fully adult specimen used for illustrating the details of the ventral valve ornamentation. (NHMW 87192 /GP 250). 4 a: detailed view of the spiny tubercles and of the very small scales on the surface. Fig. 5. Paratype (NHMW 87192 /GP 251). Articulated early juvenile specimen illustrating the ontogeny for this species. 5 a: internal view of the dorsal valve with the brachidium at early cucullate phase of development. The crura are just beginning to emerge. The commissural tuberculation of the valve is still reduced. 5 b: detailed view of the ventral valve ornamentation. The rows of tubercles and the scaly surface are already developed. 5 c: detailed ventral view of the brachidium. Note that the pedestal is already developed. 5 d: anterior view of the brachidium. 5 e: oblique lateral view of the brachidium. 5 f: detailed view of a crus in development (arrows). Fig. 6. Paratype. Articulated juvenile specimen illustrating later stage of the ontogeny (NHMW 87192 /GP 252). 6 a: internal view of the dorsal valve with the brachidium at a second stage of development. The resorption process of the posterior part of the cucullate structure is completed and claws in their first appearance are produced. 6 b: detailed view of the ventral ornamentation. 6 c: detailed anterior view of the brachidium. 6 d: detailed posterior view of the brachidium showing the resorption process when completed., Published as part of Simon, Eric, Logan, Alan, Zuschin, Martin, Mainguy, Jerome & Mottequin, Bernard, 2016, Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae, pp. 1-34 in Zootaxa 4137 (1) on page 25, DOI: 10.11646/zootaxa.4137.1.1, http://zenodo.org/record/263667

Mottequin, Bernard (2008): New observations on Upper Devonian brachiopods from the Namur-Dinant Basin (Belgium). Geodiversitas 30 (3): 455-537, DOI: http://doi.org/10.5281/zenodo.5378729, {"references":["ASSELBERGHS E. 1912. - Description d'une faune frastheir care.Marie Legrand-Blain (Gradignan) advised nienne inferieure du bord nord du bassin de Namur. on systematics of the Productidina.Wilfried Miseur Bulletin de la Societe belge de Geologie, de Paleontologie (Brussels) did the photographic work. Reviews by et d'Hydrologie, Memoires 26: 1-47. Denise Brice (Lille) and Patrick Racheboeuf (Brest) ASSELBERGHS E. 1923. - La faune de la Grauwacke de and the editorial revision by Annemarie Ohler Rouillon (base du Devonien moyen). Memoires du","Musee royal d'Histoire naturelle de Belgique 33: 1-76. (Paris) helped to improve the manuscript. George ASSELBERGHS E. 1936. - Le Devonien du bord nord Sevastopulo (Dublin) advised on linguistic aspects du Bassin de Namur. Memoires de l'Institut geologique of the paper. I am deeply grateful to all of them. de l'universite de Louvain 10: 229-327.","ASSELBERGHS E. & MAILLIEUX E. 1925. - Comptes rendus de la cinquieme session extraordinaire de la REFERENCES Societe geologique et mineralogique de Bretagne tenue dans le terrain devonien de l'Ardenne, entre ABOUSSALAM Z. S. 2003. - Das \"Taghanic-Event\" im Charleville et Gembloux du 13 au 19 avril 1925. hoheren Mitteldevon von West-Europa und Marokko. Bulletin de la Societe geologique et mineralogique de Munstersche Forschungen zur Geologie und Palaontolo- Bretagne 6: 117-187. gie 97: 1-332. BALINSKI A. 1977. - Biernatella - a new Devonian ALEKSEEVA R. E. 1969. - [New and unusual Lower double-spired brachiopod.Acta Palaeontologica Polo- Devonian spiriferids (suborder Athyrididina]. 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Genus Thomasaria Stainbrook, 1945 TYPE SPECIES. — Thomasaria altumbona Stainbrook, 1945, by monotypy., Published as part of Mottequin, Bernard, 2008, New observations on Upper Devonian brachiopods from the Namur-Dinant Basin (Belgium), pp. 455-537 in Geodiversitas 30 (3) on page 512, DOI: 10.5281/zenodo.5378729

Genus Acutatheca Stainbrook, 1945 TYPE SPECIES. — Acutatheca propria Stainbrook, 1945, by monotypy., Published as part of Mottequin, Bernard, 2008, New observations on Upper Devonian brachiopods from the Namur-Dinant Basin (Belgium), pp. 455-537 in Geodiversitas 30 (3) on page 502, DOI: 10.5281/zenodo.5378729