taxonID	type	description	language	source
ED69878E097EFB1DFF2FFC52FC5A26ED.taxon	materials_examined	TYPE: — ‘ Kuznetsk’. No diacritical marks can be used in specific epithets (McNeill et al. 2012; Art. 60) therefore Pantocsek’s original spelling armaševskii is here transcribed phonetically as armashevskii.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097EFB1DFF2FFA96FE2A27CF.taxon	materials_examined	TYPE: — ‘ Calcaire du Sendai’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097EFB1DFF2FFA75FD0D248A.taxon	materials_examined	TYPE: — ‘ Chimborazo, Springfield Estate, Mount Hillaby’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097EFB1DFF2FF931FCD1256A.taxon	materials_examined	TYPE: — ‘ Barbados’ (ANSP SchAR 3461, holotype! = Fig. 214).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097EFB1CFF2FF80FFE8A22EB.taxon	description	(LM: Fig. 212) ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 29	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097AFB18FF2FFA34FA652A47.taxon	description	(SEM: Figs 179 – 184; LM: Figs 185 – 192)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097AFB18FF2FFA34FA652A47.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, Cambridge Estate’ (BM 3072, holotype! = Fig. 188; Williams 1988: 53).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097AFB18FF2FFA34FA652A47.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, Cambridge Estate’ (BM 3044, holotype! = Fig. 192). Triceratium uviferum A. W. F. Schmidt (1885: taf. 88, fig. 14). TYPE: — ‘ Barbadoes’. Triceratium antipodum Pantocsek (1892: pl. XXXVI, fig. 514; 1905: p. 106). TYPE: — ‘ Oamaru in Nova Seelandia’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097AFB18FF2FFA34FA652A47.taxon	materials_examined	TYPE: — ‘ Clealand, Colonial, Mount Hillaby’. Frustules subrectangular in girdle view, valves tripolar, with straight (Fig. 179), slightly concave (Fig. 180), or gently convex sides (Fig. 186). Valve face strongly undulate, with a central trifolium (Fig. 181). Central parts of the projections generally depressed, valve margins raised (Fig. 181). At each pole, there is a low elevation terminated in a rounded pseudocellus (Fig. 179). An inconspicuous hyaline marginal ridge extends between each pseudocellus (Fig. 181). Additional ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 25 anastomosing hyaline ridges are located on the trifolium (Figs 179, 181). Except for a small hyaline area in the centre of the trifolium (Figs 179, 182), whole valve face is perforated by poroid areolae (Figs 185 – 192) occluded by external cribra (Fig. 182). Within the depressed parts of the projections, areolae form well-defined parallel rows, although the density of areolation sometimes varies (Figs 185 – 192). Along the valve margins and on the trifolium, areolae tend to be smaller and more densely packed (Figs 185, 189). At the junction of the trifolium sectors, there is a distinct, single large areola (Figs 179 – 180, 185, 189). Mantle is steeply downturned, shallow (Fig. 181), with a smooth hyaline margin that is slightly expanded inwardly (Fig. 181), and in some specimens conspicuously stepped (Fig. 184). Whole mantle is perforated by poroid areolae with cribra (Fig. 184). On the valve interior, along each margin between the trifolium and the pseudocellus, there is a series of short internal costae (Figs 181, 185 – 192). These costae and the expanded mantle margin provide attachment for the valvocopula, which is held by clasping devices and a fossa (Fig. 183). Valvocopula is closed, deeper than the mantle, mostly hyaline, with two separate rows of poroid areolae occluded by simple cribra (Figs 183 – 184). Other detached girdle element was observed, which may represent a copula or a pleura (Fig. 183). This element is closed, shallow, and bears a single row of poroids that are variable in size. Along the advalvar edge of this element, there is a prominent hyaline pars interior which diminishes in size next to the poles (Figs 183 – 184). Measurements (n = 10): average side length: 42.5 – 94.3 µm; 3 – 4 areolae in 10 µm (measured within the depressed sectors); 3 – 4 costae in 10 µm measured along the valve face margin. No rimoportulae were found. Geographic and stratigraphic distribution (Fig. 10, sites 14, 18): (a) specimens: Middle Eocene-early Miocene: Barbadoes: BM stubs P. 405 (Figs 179, 181) and P. 1347 (Figs 180, 182 – 184), BM 3044 (Fig. 192), BM 61079, BM coll. Adams TS 266 (Fig. 191). Oceanic Formation outcrops at Joe’s River: BM 63574 (Fig. 186), BM 65967, BM 68637 (Fig. 185); Chalky Mount: BM 61082 (Fig. 187), BM 66525 (Fig. 189); Cambridge: BM 3072 (Fig. 188, holotype); Springfield: BM 37839 (Fig. 190), BM coll. Adams F 1312. (b) records: Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Pantocsek (1892: pl. XXXVI, fig. 514; 1905: 106). Middle Eocene-early Miocene: Barbadoes: Schmidt (1885: taf. 88, fig. 14), Hustedt (1930: taf. 371, fig. 5). Oceanic Formation outcrops at Chalky Mount: Schmidt (1886 a: taf. 94, fig. 17); Cambridge: Greville (1864 b: 93, pl. XIII, fig. 15; 1866 b: 83, pl. IX, fig. 19); Clealand, Colónial and Mount Hillaby: Brun (1896: 245, pl. XXI, figs 19 – 23). Observations: — In LM, the conspicuous central opening on the trifolium may appear as a rimoportula. When carefully focusing through this structure, however, it is evident that no internal slit is associated with this large opening. This has been corroborated by SEM observations, and therefore E. foveata is one of the few species of Entogoniopsis that lack rimoportulae. Entogoniopsis foveata has a complex taxonomic history that includes many independently published names that have never been synonymized. In BM, there are type specimens for both Triceratium foveatum Greville (BM 3072; Fig. 188) and T. picturatum Greville (BM 3044; Fig. 192) (Williams 1988: 53, 57). Examination of both slides indicates no morphological differences between these taxa and therefore they are interpreted as conspecific (Figs 188, 192). Although no specimens of T. uviferum A. W. F. Schmidt (1885: taf. 88, fig. 14) nor T. antipodum Pantocsek (1892: pl. XXXVI, fig. 514) are available, they can be treated as junior synonyms of Triceratium foveatum based on the presence of raised pseudocelli and the pronounced trifolia with a large central areola in both. BM 68637, from Joe’s River, Barbadoes (Fig. 182), has a mounted specimen labelled ‘ T. lobato-gemmatum’ [sic] Brun. The morphological features of this specimen agree with Brun’s (1896: 245, pl. XXI, figs 19 – 23) description and illustrations, but also do not differ considerably from the type specimen of T. foveatum. Therefore T. lobatogemmatum is included as a synonym of the latter. In the description of T. lobatogemmatum, Brun (1896: 245) remarked that this species is ‘ closely allied’ with Triceratium westianum Greville (1861 a: 43, pl. IV, fig. 11), Triceratium figuratum Greville (1865 b: 101, pl. IX, fig. 15), Triceratium lobatum Greville (1863: 233, pl. IX, fig. 13), Triceratium nitescens Greville (1865 a: 8, pl. II, fig. 19), Triceratium antipodum Pantocsek (1892: pl. XXXVI, fig. 514; 1905: p. 106), and Triceratium tripes Brun (1896: 246, pl. XXI, fig. 14), and that all these taxa may prove to be varieties within a single species. As presented here, some of these belong to Entogoniopsis, others to Trilamina (see below). Notably, however, Brun made no reference to either T. foveatum or T. picturatum, even though the illustrations provided by Greville (1864 b: pl. XIII, fig. 15; 1866 b: pl. IX, fig. 19) fit well within the range of morphological variation documented in Brun’s line drawings (Brun 1896: pl. XXI, figs 19 – 23). Triceratium picturatum was reported as extant from Dunedin, New Zealand (Wood 1963: 196). Wood’s specimen, however, does not have internal costae and appears to lack a trifolium (Wood 1963: pl. III, fig. 48). All verified reports of E. foveata consistently indicate the middle Eocene through possibly Miocene as the stratigraphic range of this species. 26 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB16FF2FFE37FE642557.taxon	description	(SEM: Figs 161 – 167, LM: Figs 168 – 170)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB16FF2FFE37FE642557.taxon	materials_examined	TYPE (here designated): — ‘ DODO- 123 - D 1 dredging, Mascarene Ridge, Indian Ocean’ (BM 101699, holotype! = Fig. 170). Valve outline tripolar (Figs 168 – 170), with slightly rounded poles, and straight to gently convex sides (Figs 161, 165). Valve face undulate, with a pronounced trifolium that bears a broad, irregular hyaline area at its centre (Figs 161 – 162). At each pole, there is a low polar elevation, terminated in a flat hyaline plate bearing a pseudocellus (Figs 162 – 163). A distinct hyaline marginal ridge extends between each polar elevation, its crest level with the summits of the elevations (Fig. 162). The valve face-marginal ridge junction is marked with a row of large openings through the hyaline marginal ridge that follow the undulation of the valve face (Fig. 162). Additionally, the hyaline marginal ridge is supported by series of fine external costae that protrude a short distance toward the valve face centre (Figs 161 – 162). Near the margins of the central hyaline area, there is a group of rimoportulae, sometimes in the form of a poorly defined ring (Fig. 161). Rimoportulae open to the outside as long, slender tubes (Fig. 167), and to the inside as short straight slits across slightly raised papillae (Fig. 165). The density of areolation on the valve face varies: near the valve centre areolae are smaller and sparse (Fig. 161). Within the depressed sectors, areolae are evenly spaced and arranged in radial rows, locally curved (Figs 161 – 162). Mantle extremely shallow, steeply downturned, entirely hyaline, except for pseudocelli and scattered poroids on the distal faces of the polar elevations (Figs 162 – 163, 165). Mantle margin is hyaline, smooth, slighty expanded inwardly (Fig. 164). Along each side on the valve interior, there is a series of inconspicuous costae (Fig. 164). Valvocopula is closed, deeper than the mantle (Fig. 162), and hyaline, except for a single row of poroids adjacent to the fossa (Figs 165 – 166). Valvocopula attaches by means of small clasping devices that hold the internal costae, and the fossa, which clasps the inward expansion of the mantle margin (Figs 165 – 166). Measurements (n = 6): average side length: 128.5 – 182.0 µm; 4 internal costae in 10 µm; 3 external costae in 10 µm; 5 – 6 areolae in 10 µm measured in the central part of a depressed sector; 2 – 25 rimoportulae.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB16FF2FFE37FE642557.taxon	etymology	Etymology: — named in memory of the late Zoya Il’inichna Gleser (1929 – 2006), the respected Russian siliceous microfossil worker, who proposed the genus Sheshukovia. Geographic and stratigraphic distribution (Fig. 10, sites 2, 16): (a) specimens: Eocene: Indian Ocean, DODO- 123 - D 1 dredging, Mascarene Ridge: BM stub P. 1324 (Figs 161 – 167), BM 101699 (Figs 168 – 170). Early-middle Eocene: Western North Atlantic Ocean: ODP Site 1051: SZCZ 18906> 25 µm B, 21654> 25 µm D. (b) records: Locality and age unspecified: Cooper & Harvey (2003: fig. 15, as ‘ Trigonium dobreanum (Norman ex Greville) Hustedt'). Observations: — Entogoniopsis gleseri bears some resemblance to Triceratium rzehakii Pantocsek (1889: 94). Pantocsek’s illustration appears to show the presence of external costae on the valve face (Pantocsek 1892: taf. I, fig. 14). However, T. rzehakii seems to lack a trifolium, whose presence can sometimes be successfully interpreted from illustrations (see Entogoniopsis foveata, below). Instead, the valve face in T. rzehakii seems to rise toward the poles, which appear to be distinctly cut off from the valve face centre. Pantocsek (1889, 1892) recorded T. rzehakii in deposits from Veľký Krtíš (formerly Nagy-Kürtös), Slovakia, and two Moravian localities. Unfortunately, no material from any of these deposits is available, and type slide no longer exists in BP (K. Buczkó, personal communication, 2013).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0968FB0DFF2FFAEDFB9D2388.taxon	description	(SEM: Figs 24 – 30; LM: Figs 31, 33 – 36)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0968FB0DFF2FFAEDFB9D2388.taxon	materials_examined	TYPE (here designated): — ‘ Oamaru, Otago, New Zealand’ (BM 46607, holotype! = Fig. 34).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0968FB0DFF2FFAEDFB9D2388.taxon	description	ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 11 expanded inwardly (Fig. 25). On some valves, immediately above the hyaline margin, there is a circumferential furrow extending along each side of the valve (Fig. 24). Mantle is areolate; poroid areolae are arranged in rows generally parallel to the pervalvar axis, but less-well defined than rows on the valve face (Figs 24 – 25). Mantle areolae also show variable diameters. On the valve interior, there is a prominent network of robust costae that extend between the mantle margin and the margin of the depressed central area (Figs 25, 29). The internal costae define sectors for areolae both on the valve face and on the mantle (Figs 25, 31, 33 – 36). Girdle is composed of multiple closed bands. Valvocopula attaches to the internal costae by means of small clasping devices (Figs 29 – 30), and to the expanded mantle margin by means of a fossa (Fig. 29). The valvocopula is narrow, perforate, with poroid areolae arranged in rows parallel to the pervalvar axis (Fig. 29). A detached closed copula with a fringed pars interior was also observed (Fig. 29), bearing rows of poroid areolae similar to those on the valvocopula (Fig. 29). Measurements (n = 14): average side length: 68.43 – 263.85 µm; 2 – 4 coarser and 2 – 5 finer areolae in 10 µm; 2 costae in 10 µm, measured along the valve face margin; number of rimoportulae: 8 – 20; pseudocelli: 5 – 7 porelli in 10 µm. Geographic and stratigraphic distribution (Fig. 10, sites 12, 18 – 20): (a) specimens: Late Eocene: Falkland Plateau, South Atlantic: Vema Cruise 17, Core 107: BM stub P. 1275 (Figs 26, 28 – 30); South Tasman Rise, Southwestern Pacific Ocean: DSDP Site 281, Cores 281 - 14 through 281 - 16: SZCZ 15073, 21078, 22054 – 22056, 22057 (Fig. 33). Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: BM 46607 (Fig. 34, holotype), BM 63654 (Fig. 36); BM coll. Adams: J 671, G 86; Oamaru Diatomite outcrops at: Allan’s Farm: BM coll. Adams TS 426 (Fig. 31); Jackson’s Paddock: BM coll. Adams F 1348; ‘ Railway Cutting 1 ’: BM coll. Adams G 68 (Fig. 35). Early Oligocene: Falkland Plateau, South Atlantic: Vema Cruise 18, Core 112: BM stub P. 1350 (Figs 24 – 25, 27). (b) records: Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Reed (1991: pl. 17, fig. 215, as T. morlandii); Desikachary & Sreelatha (1989: 260, pl. 63, fig. 3, as T. morlandii var. aperta). Observations: — Grove & Sturt (1887 a: 8) reported a diatom they provisionally identified as Entogonia davyana (Greville) Greville (1863: 236; basionym: Triceratium davyanum Greville 1862 b: 232, pl. 10, fig. 4) from the fossil deposit in Oamaru, New Zealand. They provided no illustration and suggested that their specimens differed from E. davyana sensu stricto (Fig. 32) by the absence, in the Oamaru specimen, of “ the transverse lines [i. e., internal costae] in the central compartment, and the clear spaces at the base of each process [i. e., openings of the internal coil next to the proximal sides of polar elevations, see Fig. 4] ”. Subsequent to the series of papers published by Grove & Sturt (1886, 1887 a, b, c) on the diatoms from Oamaru, Grunow (1887 b, 1888) offered comments on numerous taxa proposed by them. With respect to Triceratium morlandii, Grunow (1888) wrote: “ Tr. Morlandii Gr. et St. var. sub. aperta Grun. Formen, bei denen der Ring von Meschen, die das Centrum umgeben, an 3 Stellen mehr oder weniger geöffnet ist. Sehr ähnlich solchen Formen ist Entogonia Davyana Gr. et St. (nec Grev.), die Ref. vorläufig als var. aperta bezeichnet. Tr. venosum kommt bisweilen mit dreieckiger, erweiterter Mitte vor, und hat dann eine gewisse Aehnlichkeit mit letzterer. Die Autoren erwähnen noch brieflich eine dem Tr. venosum verwandte Form mit convexen Seiten und gebogenen Rippen, die sie als var. flexuosa bezeichnen. Vielleicht kann man alle diese Arten zu Entogonia ziehen ” (Grunow 1888: 40 – 41; spelling as in original). Grunow’s views appeared in an English translation by G. C. Karop, with further annotations by Grove (Grunow 1889: 388). The relevant paragraph of the translation reads: “ Tr. Morlandii, Gr. and St. Herr Grunow proposes to add var. sub-aperta, Grun., for varieties in which the ring of meshes round the centre is more or less incomplete in three places. Such forms generally resemble Entogonia davyana, Gr. and St. (nec Grev.), which Herr Grunow provisionally designates var. aperta. ” Neither the German nor English text makes it clear whether Grunow was proposing his new variety for T. morlandii or E. davyana. In spite of the ambiguity in the German and English versions of Grunow’s accounts (1888, 1889), it has generally been accepted that Grunow meant Triceratium morlandii var. aperta: VanLandingham (1969; 1978) and the online Catalogue of Diatom Names (Fourtanier & Kociolek 2011) list T. 12 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. morlandii var. aperta rather than E. davyana var. aperta. Additionally, the Catalogue of Diatom Names identifies the name T. morlandii var. aperta as invalid but no discussion of its taxonomic status is evident. Regardless of these nomenclatural issues, our observations indicate that Entogoniopsis morlandii sensu stricto differs from what we identify as Grunow’s var. aperta, and therefore the latter should be recognised as a species. Since the full name of the new variety was not explicitly stated by Grunow (1888, 1889), the name ‘ T. morlandii var. aperta ’, as far as we can tell, has never been validly published. As no valid name exists, these specimens constitute a new species, for which the name Entogoniopsis grunowii is proposed here.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0972FB10FF2FFE7FFDFF23D0.taxon	description	(SEM: Figs 87 – 93; LM: Figs 94 – 96)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0972FB10FF2FFE7FFDFF23D0.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, from Cambridge Estate’ (BM 2873, holotype! = Fig. 94; Williams 1988: 54).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0972FB10FF2FFE7FFDFF23D0.taxon	materials_examined	TYPE: — ‘ Barbados’ (SchAR 3460, holotype! = Fig. 96; Boyer 1922: 4). Valves tripolar to subcircular, with slightly to strongly convex sides and acute poles (Figs 94 – 96). Valve face slightly depressed in the centre, and flat along the margins (Figs 87 – 88). Whole valve is perforated by poroid areolae: within the central depression, the areolae are conspicuously larger, more distantly spaced, and arranged in poorly defined rows (Figs 87 – 88). Coarse areolae are conspicuously domed (Fig. 93). The flat, marginal zone of the valve face, and the mantle are perforated by areolae that are not domed (Fig. 92), more closely spaced, and arranged in parallel rows (Figs 87 – 88). Both types of areolae are occluded by cribra (Figs 92 – 93). Mantle steeply downturned, relatively shallow, sometimes stepped (Fig. 91), with a smooth, hyaline margin that is slightly expanded outwardly and inwardly (Figs 87, 89, 91). The mantle bears numerous furrows. A prominent, circumferential furrow is located immediately above the expanded margin of the mantle (Fig. 87). Shorter furrows, parallel to the pervalvar axis (Figs 87, 91), are associated with the internal costae that span the whole depth of the mantle, and reach to the margin of the central depression on the valve face (Figs 89 – 90). These costae, together with the expanded mantle margin, provide attachment for the valvocopula, which attaches by means of small clasping devices (Fig. 90) and a fossa (Fig. 89). Valvocopula is closed, comparable in depth to the mantle, and mostly hyaline, with only two rows of poroids (Figs 88 – 90). Measurements (n = 5): average side length: 75.4 – 124.0 µm; 2 – 3 areolae in 10 µm in the central depression; 2 costae in 10 µm, measured along the valve face margin. Geographic and stratigraphic distribution (Fig. 10, sites 2, 14, 27): (a) specimens: Middle Eocene: ODP Site 1051: SZCZ 16096 B, 17945 B (Fig. 95). Middle Eocene-early Miocene: Barbadoes: ANSP SchAR 3460 (Fig. 96); Oceanic Formation outcrop at Cambridge Estate: BM 2873 (Fig. 94, holotype). Middle-late Miocene: San Clemente: BM stub P. 1346 (Figs 87 – 93). (b) records: Middle Eocene: ODP Site 1051, Cores 1051 A- 8 H and 1051 B- 9 H: (Witkowski et al. 2014).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0972FB10FF2FFE7FFDFF23D0.taxon	description	ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 17 Entogoniopsis. Valve face exterior in Stictodiscus often displays an ornamentation composed of fine, hyaline ridges (see fig. D on p. 217 in Round et al. 1990), which are absent in Entogoniopsis. Finally, although some Stictodiscus spp. possess internal costae, they are not as prominent as in E. inflata, and therefore it seems unlikely that they provide valvocopula attachment. Because of the characteristic, inflated valve outline, numerous specimens in BM (e. g., BM 61113, BM 61114, BM coll. Adams TS 266) have been identified as Triceratium inflatum Greville. On closer examination, however, the specimens on these slides are distinct from E. inflata, as they have central rimoportulae. These specimens (Figs 145 – 146), which are similar to Triceratium turgidum A. W. F. Schmidt (1882: taf. 77, fig. 18), are discussed under Entogoniopsis lineolata (below).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB19FF2FF806FC462783.taxon	description	(SEM: Figs 171 – 176, LM: Figs 177 – 178) 24 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB19FF2FF806FC462783.taxon	materials_examined	TYPE (here designated): — ‘ DODO- 123 - D 1 dredging, Mascarene Ridge, Indian Ocean’ (BM 101700, holotype! = Fig. 177). Valve outline tripolar (Figs 177 – 178), with acute or slightly rounded poles, and convex sides (Figs 171 – 172, 177 – 178). Valve face undulate, with a pronounced trifolium and deeply depressed sectors (Fig. 171). At each pole, there is a low, stout elevation, whose summit is flattened, with a small pseudocellus located on the distal face (Fig. 175). The flattened parts of the polar elevations are cut off from the remainder of the valve face by a hyaline marginal ridge extending across the maximum deflection of each elevation (Fig. 175), and between the elevations (Fig. 171). At the centre of the trifolium, there is a group of rimoportulae, which sometimes are arranged in an irregular ring (Figs 171 – 174). The external openings of the rimoportulae are in the form of simple pores or short tubes (Fig. 173), and the internal openings are barely raised slits (Fig. 174). Whole valve face is perforated by poroid areolae arranged radially in fascicles, with one fascicle per each depressed and raised sector (Figs 173, 177 – 178). On the raised sectors of the trifolium, some areolae tend to be elongated (Fig. 173). Mantle is steeply downturned, shallow (Figs 171 – 172), perforated by poroid areolae arranged in rows that are parallel to the pervalvar axis close to the mid-points of each side, but become divergent closer to the poles (Fig. 171). Mantle margin is smooth, hyaline, and slightly expanded both inwardly and outwardly (Figs 171, 176). Along each side on the valve interior, there is a series of robust costae that arise from the mantle margin and protrude a short distance toward the valve face centre (Figs 176 – 178). Measurements (n = 5): average side length: 84.0 – 196.3 µm; 1 internal costa in 10 µm; 2.5 – 3 areolae in 10 µm measured in the central part of a depressed sector; 3 – 7 rimoportulae.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0975FB19FF2FF806FC462783.taxon	etymology	Etymology: — named for Klaus Kemp, who has lent us his experience in diatom cleaning and mounting on numerous occasions. Geographic and stratigraphic distribution (Fig. 10, site 16): Eocene: Indian Ocean: DODO- 123 - D 1 dredging, Mascarene Ridge: BM stub P. 1322 (Figs 171 – 176), BM 101700 (Figs 177 – 178). Observations: — Although no girdle elements were found, there is a series of internal costae present along each side, arising from an inwardly expanded, smooth and hyaline mantle margin, strongly suggesting that the valvocopula attaches by means of a series of clasping devices and a fossa. Also the presence of a pronounced trifolium supports placement in Entogoniopsis. Those specimens of Entogoniopsis kempii that have uniformly curved sides (e. g., Fig. 172) resemble Entogoniopsis lahusenii comb. nov. (see below) in valve outline and the arrangement of internal costae. However, as figured in Pantocsek (1889: pl. VII, fig. 124), E. lahusenii appears to lack the central trifolium. The valve outline of Entogoniopsis saratoviana comb. nov. is also similar to that of E. kempii, especially the specimens with angular convexities on the sides (e. g., Fig. 171). The figure in Pantocsek (1889: pl. VII, fig. 122), however, indicates that E. saratoviana has internal costae only at the margins of the prominent trifolium, which distinguishes it from E. kempii (e. g., Fig. 172). The polar elevations in both E. lahusenii and E. saratoviana do not appear to be cut off from the remainder of the valve face, which differs them from E. kempii. The valve outline similarities may indicate that E. kempii and either E. lahusenii or E. saratoviana represent dissimilar valves of a heterovalvar species. Until intact frustules have been examined, we retain all three as separate taxa.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFE3CFD3B208F.taxon	materials_examined	TYPE: — ‘ Kusnetzk’. Trigonium lahuseni (Pantocsek) Hustedt (1959 a: taf. 468, fig. 9).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0976FB14FF2FFEE5FB5B20F2.taxon	description	(SEM: Figs 128 – 141; LM: Figs 142 – 144, see also Figs 145 – 148)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0976FB14FF2FFEE5FB5B20F2.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, from Cambridge estate’ (BM 2743, holotype! = Fig. 142; see Williams 1988: 55).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0976FB14FF2FFEE5FB5B20F2.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, Cambridge estate’ (BM 3263, syntype! = Fig. 143; see Williams 1988: 58). Triceratium turgidum A. W. F. Schmidt (1882: taf. 77, fig. 18). TYPE: — ‘ Barbadoes’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0976FB14FF2FFEE5FB5B20F2.taxon	materials_examined	TYPE: — ‘ Barbados’ (ANSP SchAR 3459, holotype! = Fig. 144). Trigonium quadratum (Greville) Hustedt (1959 a: taf. 470, figs 7 – 8). Valves tri-or quadripolar in valve view, with high, slender polar elevations bearing small, hemispherical pseudocelli at the summits (Figs 128 – 129, 136, 139). The valve face is gently depressed next to the marginal hyaline ridge in some specimens (Fig. 130), but always conspicuously domed at the centre (Figs 130, 136). On the central dome, there is an irregular ring of rimoportulae (Figs 133 – 134, 140 – 141). External openings of the rimoportulae in the form of short, non-buttressed tubes (Fig. 133). Internal openings as slightly raised slits (Figs 134, 141). Poroid areolae are aligned in radial rows, and generally increase in size towards the valve face margin (Figs 128, 136), excepting few areolae scattered within the ring of rimoportulae. Mantle steeply downturned, relatively shallow (Figs 135, 139), with a hyaline margin, slightly expanded toward the interior (Fig. 132), and sometimes also toward the exterior (Fig. 139). Mantle areolate, with rows of areolae parallel to the pervalvar axis near the mid-point of each side, becoming divergent next to the valve poles (Figs 130, 136). The mantle areolae adjacent to the marginal hyaline ridge sometimes tend to be elongate (Fig. 130). Small tubercles are scattered on the distal face of the hyaline ridge adjacent to the mantle (Figs 135, 139). Numerous spinules are arranged in more or less well-defined rings on the valve face (Figs 133, 139), and parallel rows of granules are present on the mantle (Figs 135, 139). Short internal costae are distributed in irregular intervals along each side (Figs 131, 137), reaching only a short distance toward the valve face centre (Figs 132, 137 – 138). Measurements (n = 10): average side length: 50.3 – 134.7 µm; 1 costa in 10 µm measured along the valve face margin; 3 areolae in 10 µm measured near the valve face margin; 2 – 10 rimoportulae. Geographic and stratigraphic distribution (Fig. 10, sites 8, 14, 16; records of T. turgidum below are marked with an asterisk): (a) specimens: Middle Eocene: North Atlantic Ocean: DSDP Site 6, Bermuda Rise: BM stub P. 1353 (Figs 136 – 137, 139 – 141). Middle-late Eocene transition: Indian Ocean: DODO- 123 - D 1 dredging, Mascarene Ridge: BM stub P. 174 (Figs 128 – 135). Middle Eocene-early Miocene: Barbadoes: BM coll. Adams: GC 3447 * (Fig. 148), TS 266 *; SchAR 3459 (Fig. 144); Oceanic Formation outcrops at Cambridge Estate: BM 2743 (Fig. 142, holotype), BM 3263 (Fig. 143), BM coll. Adams F 1403 (Fig. 145); Chalky Mount: BM 61114 *; Joe’s River: BM stub P. 1351 (Fig. 138), BM coll. Adams TS 929 (Fig. 147); Springfield: BM 61113 *, BM coll. Adams: GC 3445 *, GC 3448 * (Fig. 146); Newcastle: BM coll. Adams GC 3446 *. (b) records: Middle Eocene-early Miocene: Barbadoes: Boyer (1922: 4, pl. I, fig. 5), Schmidt (1882: taf. 77, fig. 18 *), Hustedt (1959 a: taf. 470, figs 7 – 8); Oceanic Formation outcrops at Cambridge Estate: Greville (1863: 233, pl. X, fig. 16; 1865 b: 104, pl. IX, fig. 19). Observations: — Beside the difference in number of poles, Triceratium lineolatum Greville and Triceratium quadratum Greville are morphologically indistinguishable, suggesting that they are tri-and quadripolar variants of the same species; Greville identified both from the same locality, Cambridge Estate (Greville 1863: 233, Greville 1865 b: 104). Williams (1988: 58) indicated that in the protologue for T. quadratum, BM 3263 is given as a slide number only, and illustrated as a syntype a fractured specimen from BM 2902 (Williams 1988: pl. 70, figs 2 – 3), which is annotated with MF coordinates. In this study, we located a complete specimen on slide BM 3263 (see Fig. 143), and consider this the second syntype of T. quadratum. ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 21 The specimens of E. lineolata reported from the Indian Ocean resemble Greville’s holotype specimen (Fig. 142) as both have the same areolation pattern and valve outline. However, a number of tripolar specimens have been examined from various Barbadoes deposits, which resemble E. lineolata but differ in valve outline. These specimens (Figs 145 – 146), with distinctly convex central part of each side, match the image of Triceratium turgidum (Schmidt 1882: taf. 77, fig. 18), thus we consider T. turgidum a synonym of E. lineolata under the assumption that this taxon displays considerable variation in valve outline; some slides in BM that contain specimens resembling T. turgidum have been mislabelled as E. inflata because of the valve outline (BM 61113, BM 61114, BM Coll. Adams TS 266; see discussion under E. inflata, above). Another species that we considered is Biddulphia quadrans Boyer (1922: 4). Boyer (1922) did not comment on any similarity between B. quadrans (Fig. 144, holotype specimen) and T. quadratum. Instead, he suggested a close relationship with Entogoniopsis dutertrei (= Triceratium dutertrei Pantocsek & Tempère in Pantocsek 1889: 90). Boyer may have considered B. quadrans as distinct from T. quadratum because of the slightly different valve outline: Greville’s line drawing of T. quadratum shows a precisely quadrate specimen (Greville 1865 b: pl. IX, fig. 19) but the syntype of T. quadratum has slightly concave sides and strongly resembles B. quadrans (Fig. 143), further confirming the valve outline variability. The holotype specimen of B. quadrans (Fig. 144) is indistinguishable from the quadripolar specimens of T. turgidum (Fig. 148), which lends further support to our hypothesis that E. lineolata is synonymous with T. quadratum, B. quadrans and T. turgidum.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0973FB13FF2FFE7FFDEA2096.taxon	description	et stat. nov. (SEM: Figs 97 – 101; LM: Figs 102 – 107)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0973FB13FF2FFE7FFDEA2096.taxon	materials_examined	TYPE (here designated): — ‘ Oamaru, Otago, New Zealand’ (BM 46554, holotype! = Fig. 105).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0973FB13FF2FFE7FFDEA2096.taxon	description	Typification: — The first mention of Triceratium venulosum (not identified as a variety) in Sturt’s unpublished catalogue is on p. 134, slide BM 46554. This specimen is in valve view, with a minor portion of the valve face obscured by debris, and has girdle elements attached (Fig. 105), which matches the specimen figured in Grove & Sturt (1886: pl. XIX, fig. 15). Also its size agrees well with the published illustration. Other specimens indicated in Sturt’s catalogue are either larger (BM 46556), or situated with the valve tilted (BM 46575). The specimen on slide BM 46554, however, is not as densely areolated as the one illustrated by Grove & Sturt (1886); it is possible that the areolation pattern on the figure was modified to show what the authors considered typical morphology. Therefore we conclude that the specimen on slide BM 46554 is the holotype of Entogoniopsis major (Fig. 105). Geographic and stratigraphic distribution (Fig. 10, site 18): (a) specimens: Late Eocene-earliest Oligocene: Oamaru, New Zealand: BM stub P. 1337 (Figs 97 – 101), BM coll. Adams: G 3, G 28 (Fig. 107), G 100, G 659, GC 3304, BM 46554 (Fig. 105, holotype), BM 61142, BM 61276. Oamaru Diatomite outcrops at: ‘ Railway Cutting 9 ’: BM coll. Adams G 72; Allan’s Farm: BM coll. Adams G 89, G 614 (Figs 102, 104); Forrester’s: BM 37914 (Fig. 106), BM 37916 (Fig. 103); Jackson’s Paddock: BM coll. Adams G 116; Cormack’s Siding: BM 61141. Locality and age unspecified: BM coll. Adams TS 317. (b) records: Late Eocene-earliest Oligocene: Oamaru, New Zealand: Schmidt (1888: taf. 127, figs 1 – 2, as T. majus, and fig. 3, as T. venulosum; 1891: taf. 168, fig. 5, as T. majus). Oamaru Diatomite outcrop at Cormack’s Siding: Grove & Sturt (1886: 327, pl. XIX, figs 15 – 16), De Lautour (1888: 311, pl. XXI, fig. 1). 18 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. Observations: — The original publication of this species as a variety of Triceratium venulosum Greville (1864 b: 90) has caused considerable confusion. Grove & Sturt (1886: 328) distinguished the variety based on the diameter of the valves, which are much larger in T. venulosum var. major than in Greville’s species. Grunow (1888) suggested the replacement species name T. majus for T. venulosum var. major and Grove’s later annotations (in Grunow 1889) indicate that Grove & Sturt accepted this name. Yet, Greville’s figure of T. venulosum (1864 b: pl. XIII, fig. 21) is difficult to interpret and does not allow precise identification; furthermore, the holotype specimen of T. venulosum has not been located (Williams 1988: 60), further complicating identification. As a result, the diatom reported from the Oamaru Diatomite by Grove & Sturt (1886) as T. venulosum var. major is commonly identified as both T. venulosum Greville and T. majus Grove & Sturt. Our study shows that Entogoniopsis major is absent from Greville’s Barbadean localities, suggesting that T. venulosum is not the same as T. venulosum var. major. Finally, Simonsen, in the 1968 reprinting of some of Greville’s papers, synonymized T. venulosum with Triceratium tabellarium Brightwell (1856: 275), although he gave no reasons for doing so (Simonsen 1968: xii; see discussion on Entogoniopsis tabellaria, below). In summary, T. venulosum is not the same as T. venulosum var. major and therefore we propose Entogoniopsis major comb. nov. et stat. nov. based on Triceratium venulosum var. major Grove & Sturt (1886: 327).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096BFB0BFF2FFDACFDBA273E.taxon	description	comb. nov. (SEM: Figs 11 – 16; LM: Figs 17 – 23)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096BFB0BFF2FFDACFDBA273E.taxon	materials_examined	TYPE (here designated): — ‘ Oamaru, Otago, New Zealand’ (BM 46565, lectotype! = Fig. 18).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096BFB0BFF2FFDACFDBA273E.taxon	description	Typification: Specimens of Triceratium morlandii are found on several slides in the Sturt collection at the BM. A specimen on strewn slide BM 46548, annotated with Maltwood Finder (MF) coordinates 36 | 17.5, is described in the unpublished Sturt catalogue as “ perfect ” (p. 131). This specimen is considerably tilted, however, and has strongly concave sides, and therefore it is unlikely to have served as the basis for the remarkably detailed illustration in Grove & Sturt (1887 a: pl. II, fig. 5). Instead, a straight-sided, selected specimen mounted on slide BM 46565 (Fig. 18), also from the Sturt collection, is designated here as lectotype of E. morlandii. Geographic and stratigraphic distribution (Fig. 10, sites 12 – 13, 18 – 19): (a) specimens: Middle Eocene: Falkland Plateau, South Atlantic Ocean: Vema Cruise 17, Core 107: BM stub P. 1275 (Figs 12, 14). Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: BM stub P. 1277 (Figs 11, 13, 15 – 16); BM 46565 (Fig. 18, lectotype), BM 46600, BM 61155, BM 61157, BM 63654, BM 66534, BM coll. Adams: F 1347, G 3, G 84, G 86, G 93 (Fig. 22), G 133, G 142 (Fig. 20), TS 270 (Figs 21, 23), TS 291, TS 315 (Fig. 17). Oamaru Diatomite outcrops at: Allan’s Farm: BM coll. Adams TS 429; Jackson’s Paddock: BM 61159, BM 63650, BM coll. Adams G 157, G 162 (Fig. 19); Totara: BM coll. Adams TS 435; ‘ Railway Cutting’: BM coll. Adams: G 68, G 69, G 73. 10 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. (b) records:	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096BFB0BFF2FFDACFDBA273E.taxon	description	Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Laporte & Lefébure (1929: pl. 8, fig. 54), Schmidt (1888: taf. 128, figs 5 – 6; 1890: taf. 150, fig. 22); Oamaru Diatomite outcrops at: Allan’s Farm: Doig (1991: 112); Bain’s Lower, Bain’s Farm, Cormack’s Top, Forrester’s Hill: Desikachary & Sreelatha (1989: 260); Jackson’s Paddock: Desikachary & Sreelatha (1989: 260), Doig (1991: 114); Cormack’s Siding: Grove & Sturt (1887 a: 7, pl. II, fig. 5), De Lautour (1888: 310, pl. XXI, fig. 6), Desikachary & Sreelatha (1989: 260); Totara: Desikachary & Sreelatha (1989: 260); Flume Gully: Doig (1991: 125); Taylor’s Quarry: Doig (1991: 113); William’s Bluff: Desikachary & Sreelatha (1989: 260), Doig (1991: 114); Division Hill: Desikachary & Sreelatha (1989: 260), Doig (1991: 119); Mavor’s Farm: Desikachary & Sreelatha (1989: 260), Doig (1991: 120); Table Top Hill: Doig (1991: 125). Observations: — Morphological variation in Entogoniopsis morlandii is reflected primarily in the presence or absence of the central of rimoportulae, suggesting that E. morlandii may have formed heterovalvar frustules. Unfortunately, the only frustules available for study were in LM in girdle view (Fig. 23). Therefore, heterovalvy could not be directly investigated. Specimens without rimoportulae, however, tend to be of a relatively small size (e. g., Figs 17 and 19 versus Figs 18, 20 – 22). Thus, it is also possible that at some stages in the vegetative phase of the life cycle, E. morlandii produced valves without rimoportulae. Entogoniopsis morlandii has a highly variable areolation pattern, associated with the network of internal costae. Next to the midpoint of each side, the pattern of internal costae is usually uninterrupted: costae extend toward the valve face centre over a similar distance, approximately half the distance from the valve face centre to the mid-point of a side, and form sectors, in which there are usually two rows of areolae. However, closer to the poles, the costae on the valve face become shorter, and often bifurcate or intersect at variable angles. As a result, in addition to the central depression, where the areole are larger and arranged more loosely than in the marginal sectors, there are also three areas next to the proximal side of each polar elevation, in which areolae are larger and scattered (Figs 11, 17 – 22). These areas appear distinctly thickened in LM, and in some specimens are separated from the central depression by a narrow zone of particularly complex latticework of internal costae (e. g., Fig. 18). This is probably what Grunow (1889: 388) meant, when suggesting that forms „ in which the ring of meshes round the centre is more or less incomplete in three places ” should be grouped in Triceratium morlandii var. subaperta. The degree of separation between the central depression and the areas adjacent to the proximal sides of the polar elevations seems to be size-dependent (Figs 17 – 22), with none or less separation visible in smaller specimens (e. g., Fig. 21). We therefore suggest that the variation with respect to this morphological feature results from purely mechanical reasons. For further comments on the varieties of T. morlandii suggested by Grunow, see discussion under Entogoniopsis grunowii, below.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096CFB11FF2FF8CDFCE923D0.taxon	description	comb. nov. et stat. nov. (SEM: Figs 69 – 75; LM: Figs 76 – 81, see also Figs 85 – 86) ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 15	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096CFB11FF2FF8CDFCE923D0.taxon	materials_examined	TYPE (here designated): — ‘ Oamaru, Otago, New Zealand’ (BM 71533, lectotype! = Fig. 77). Triceratium Novae Seelandiae (Grove & Sturt) A. W. F. Schmidt (1888: taf. 127, figs 5 – 8).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096CFB11FF2FF8CDFCE923D0.taxon	description	Typification: A strewn slide in the Saxton collection (BM 71533), prepared by Grove and annotated with MF coordinates (31 | 14), includes a specimen of E. novazealandica in girdle view (Fig. 77). The slide label reads ‘ T. dobr. F. V. ’ (i. e., girdle view). Based on the high overall resemblance of this specimen to the one illustrated in Grove & Sturt (1886: pl. XIX, fig. 18) and reproduced here (Fig. 86), we designate slide BM 71533 as the lectotype of E. novazealandica. Geographic and stratigraphic distribution (Fig. 10, sites 18 – 19): (a) specimens: Late Eocene: South Tasman Rise, Southwestern Pacific Ocean: DSDP Site 281, core-section 281 - 15: SZCZ 15073, 22056 – 22057. Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: BM stubs P. 1274 (Figs 70, 72 – 75) and P. 1277 (Figs 69, 71), BM 46550, BM 46561, BM 46575 (Fig. 81), BM 46607 (Fig. 78), BM 61050, BM 63524 (Fig. 76), BM 71533 (Fig. 77, lectotype), BM coll. Adams: F 1295, G 93, G 659 (Fig. 80). Oamaru Diatomite outcrop at Jackson’s Paddock: BM coll. Adams G 116 (Fig. 79). (b) records: Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Schmidt (1888: taf. 127, figs 5 – 8, 1891: taf. 168, fig. 2); Laporte & Lefébure (1929: pl. XIV, fig. 99); Hustedt (1959 a: taf. 470, figs 2, 4 – 6); Oamaru Diatomite outcrops at Cormack’s Siding: Grove & Sturt (1886: 328, pl. XIX, figs 17 – 18), De Lautour (1888: 310, pl. XXI, fig. 4), Desikachary & Sreelatha (1989: 274); Bain’s Farm, Bain’s Lower, Cormack’s Top, Division Hill, Forrester’s Hill, Jackson’s Paddock, Mavor’s, William’s Bluff: Desikachary & Sreelatha (1989: 274 – 275). Observations: — Entogoniopsis novazealandica has a complicated nomenclatural history including many orthographic variations as well as an evolving taxonomic concept with respect to biddulphioid diatoms. Grove & Sturt tentatively proposed their specimens to be a variety of Triceratium dobreeanum Norman ex Greville (1865 a: 6, pl. II, figs 23 – 24, reproduced here in Figs. 82 – 83; Grove & Sturt 1886: 328, pl. XIX, figs 17 – 18, reproduced here in Figs 85 – 86). Grove & Sturt (1886) remarked that their specimens were more densely areolated and had lower polar elevations than Greville’s T. dobreeanum. The line drawing in Greville (1865 a), however, reveals another significant difference: the presence of broken linking spines at the summits of the polar elevations in T. dobreeanum. Examination of the holotype specimen of T. dobreeanum (BM 3340; Fig. 84) reveals a fractured valve with an attached valvocopula, mounted in girdle view. Both the structure of the valvocopula and the valve resemble 16 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. E. novazealandica. However, the holotype confirms the presence of damaged linking spines at the summits of the slender polar elevations, alongside the pseudocelli. Species in Entogoniopsis generally lack linking spines, except for one unnamed species (reported here as Entogoniopsis? sp. 1, below). The considerable variability in valve face undulations observed in E. novazealandica suggests that this species may have been heterovalvar. Based on the currently available evidence, however, the heterovalvy with respect to the presence or absence of linking spines cannot be verified. Finally, Greville (1865 a) reported T. dobreeanum from dredged material of unknown age, and therefore we cannot base the separation between the two taxa in question based on age difference. Thus Triceratium dobreeanum var. novazealandica is recognised as the species E. novazealandica, although future studies may demonstrate that both taxa are synonymous.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFD3CFCF3219A.taxon	materials_examined	TYPE: — ‘ Oamaru, Otago, New Zealand’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	description	Williams, comb. nov. (SEM: Figs 37 – 43; LM: Figs 44 – 49, see also Figs 50 – 51)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	TYPE: — “ Kusnetzk ” (= Entogonia saratoviana Pantocsek).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	TYPE: — “ Küsnetzk (fossile) Rare ”.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	TYPE: — “ Küsnetzk (fossile) Rare ”.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	TYPE: — “ Heiligenhafen an der Ostsee ” (BRM Zt 1 / 18, holotype = Simonsen 1987: 184, pl. 294, fig. 1).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	TYPE: — Oamaru Diatomite outcrop at Forrester’s, New Zealand (CHR 617491, holotype! of Triceratium forresterii Reed = Fig. 51).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	description	Geographic and stratigraphic distribution (Fig. 10, sites 1 – 7, 10 – 11, 14, 16 – 18): (a) specimens: Late Palaeocene: Kusnetzk, Penza District, Russia: BM 61201 (Fig. 49), BM 63960, BM coll. Adams: GC 3368, M 89. Early Eocene: Kamyshlov, Sverdlovsk District, Russia: BM stub P. 1317 (Figs 37, 39 – 43); BM 63691 (Fig. 47), BM 63692 (Fig. 46), BM 66536 (Fig. 44), BM 78395; BM coll. Adams TS 748 (Fig. 45); Kirgizskoe diatomite bed, northwestern Kazakhstan: LECB stub 14 (Fig. 38). Middle-late Eocene transition: Mascarene Ridge, Indian Ocean, dredging DODO- 123 - D 1: CAS 476021. Late Eocene-earliest Oligocene: Oamaru Diatomite outcrop at Forrester’s, New Zealand: Reed (1946; CHR 617491, Fig. 51). Middle Eocene-early Miocene: Barbadoes: BM 63693 (Fig. 48). ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 13 (b) records: Late Campanian (Late Crateceous): Ust’ Man’ya, Severnaya Sos’va river, Tyumen’sk District, Pre-Polar Urals, Russia: Strelnikova (1974: 90, pl. XXXVII, figs 1 a – d). Late Palaeocene: Kusnetzk, Penza District, Russia: Pantocsek (1889: 97, pl. VI, fig. 105), Brun (1891: 47, pl. XVI, figs. 14 – 15). Early Eocene: Kamyshlov, Sverdlovsk District, Russia: Hustedt (1959 b: taf. 478, figs 23 – 24); ‘ Eastern Slopes of Urals’, Berezovo, Khanty-Mansi Autonomous District, Russia: Krotov & Schibkova (1959: 118, figs 2.4 – 2.5). Early-middle Eocene: North Atlantic Ocean: DSDP Site 552, Core 552 - 10: Baldauf (1984: pl. 13, fig. 9).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	description	Late Eocene: Heiligenhafen an der Ostsee: Wetzel (1935: 73, pl. 8, fig. 13), Hustedt (1959 b: taf. 478, fig. 22); Western Siberia: Gleser et al. (1974: pl. XXIII, fig. 7).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096EFB0CFF2FFE37FE7D2733.taxon	materials_examined	Observations: — Triceratium polycistinorum was originally proposed by Pantocsek (1889) under the name Entogonia saratoviana from Kusnetzk. Earlier, a morphologically distinct species named Triceratium saratovianum Pantocsek (1889: 94, pl. VII, fig. 122) was published precluding the transfer of E. saratoviana to Triceratium. Thus, Pantocsek proposed the replacement name Triceratium polycistinorum Pantocsek (in Tempère 1892: 119). Although we have not examined type material for E. saratoviana or T. saratovianum (Pantocsek 1889: 94), the line drawing of Triceratium saratovianum in Pantocsek (1889: pl. VII, fig. 122) shows distinct costae, presumably located on the valve interior, and a trifolium in the central part of the valve strongly suggesting that T. saratovianum should be transferred to Entogoniopsis. This action is taken below. Brun (1891) also based his species Triceratium globulifer (and the variety rotunda) on material from Kusnetzk. Considering the range of morphological variation documented here, Brun’s diatom is indistinguishable from Entogonia saratoviana Pantocsek. However, Brun’s taxonomic concept was relatively narrow: he considered the strongly produced poles and convexity of valve sides greater than that of Pantocsek’s species and took these to be a character supporting their separation. Nonetheless, Brun (1891: 47) commented on E. saratoviana that it cannot belong in Entogonia, and that it is most likely related to T. venosum Brightwell, with which we concur. We have examined the holotype of Triceratium reedii (Reed) Desikachary & Sreelatha (1989), originally published as T. forresterii by Reed (1946). Reed (1991) acknowledged that this species is a synonym of T. polycistinorum, and this view is supported here. Notably, however, Reed (1946) is the only record of Entogoniopsis polycistinora from southern mid-latitudes (Fig. 10, site 18), and it extends the verified stratigraphic range of this species from the late Campanian to the Eocene-Oligocene transition, which is considerably longer than all other species of this genus.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFC46FC5B26C8.taxon	description	(LM: Fig. 213)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFC46FC5B26C8.taxon	materials_examined	TYPE: — ‘ Oamaru, Otago, New Zealand’. Trigonium pseudonervatum (Grove & Sturt) Hustedt (1959 a: taf. 472, fig. 8).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFB6FFE8A278B.taxon	materials_examined	TYPE: — ‘ Kusnetzk’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	description	(SEM: Figs 108 – 113; LM: Figs 114 – 116; see also Figs 117 – 119)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	materials_examined	TYPE: — ‘ Kusnetzk’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	materials_examined	TYPE: — ‘ Kusnetzk’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	materials_examined	TYPE: — ‘ Kusnetzk’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	materials_examined	TYPE: — ‘ Chimborazo; Springfield Estate; Mount Hillaby; rare’. Valves tripolar, with gently convex sides, and broadly rounded poles (Figs 114 – 116). Entire valve face deeply depressed, except for the slightly raised polar elevations with flat summits that bear prominent pseudocelli (Fig. 108). A hyaline marginal ridge extends between each pseudocellus, level with the summits of polar elevations (Figs 108, 110). Whole valve face is perforated by poroid areolae that are arranged in radial rows grouped in fascicles (Figs 108, 112). Areolae are sparser in a small, irregular area at the valve centre than over the remainder of the valve face (Figs 108 – 109, 114, 116). Occasional spinules are scattered among the areolae (Fig. 108). Mantle is steeply downturned (Fig. 110). The bottom of the central depression is not level with the hyaline mantle margin, slightly expanded inwards (Fig. 111). Mantle areolae poroid, arranged in rows parallel to the pervalvar axis (Fig. 110). Closer to the poles, rows can become slightly divergent (Figs 108 – 110). Series of short, robust costae are present on the valve interior, along each side of the valve (Figs 109, 111). The proximal terminations of the costae, in the marginal part of the depressed valve face, are connected by a narrow hyaline band, with a small hyaline arc between each pair of neighbouring costae (Figs 109, 111). Valvocopula attaches to the internal costae by means of arrowhead-shaped clasping devices (Figs 112 – 113), and to the expanded mantle margin, by means of a fossa (Fig. 113). Valvocopula closed, its depth comparable to the mantle, perforated by rows of sparse poroid areolae parallel to the pervalvar axis (Figs 110, 112). Measurements (n = 11): average side length: 45.5 – 326.1 µm; 2 – 5 areolae in 10 µm; 2 – 3 costae in 10 µm, measured along the valve face margin. Geographic and stratigraphic distribution (Fig. 10, sites 2 – 3, 14, 16): (a) specimens: Late Palaeocene: Kusnetzk, Penza District, Russia: BM 61264, BM 63786 (label reads: T. tschestnovii), BM coll. Adams H 335 (Fig. 114). Middle Eocene: ODP Site 1051, Cores 1051 B- 8 H and 1051 B- 12 H: SZCZ 18849 A, 18851 C. Middle-late Eocene transition: Mascarene Ridge, Indian Ocean: dredging DODO- 123 - D 1: BM stub P. 1322 (Figs 108 – 113), BM 101699 (Fig. 116). Middle Eocene-early Miocene: Barbadoes: BM coll. Adams GC 3416 (Fig. 115). (b) records: Late Palaeocene: Kusnetzk, Penza District, Russia (Pantocsek 1889: 95, 97, pl. II, fig. 24, pl. IX, fig. 164, pl. XIII, fig. 223).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	description	Phytotaxa 209 (1) © 2015 Magnolia Press • 19	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0970FB12FF2FFD44FEDB267E.taxon	materials_examined	Observations: — Pantocsek (1889) proposed three species, that upon closer examination of Kusnetzk material and Pantocsek’s illustrations, appear to be conspecific: Triceratium squamatum (Pantocsek 1889: 95, reproduced here as Fig. 118), Entogonia truanii (Pantoscek 1889: 97, reproduced as Fig. 119) and E. tschestnovii (Pantoscek 1889: 97, reproduced as Fig. 117). The Latin diagnoses provided by Pantocsek (1889: 95, 97) indicate that the valves of both E. tschestnovii and E. truanii are convex, while there is no explicit reference to valve convexity of T. squamatum. Furthermore, T. squamatum is supposed to have more finely areolated valves, with 4 – 5 areolae in 10 µm, while E. truanii and E. tschestnovii can be distinguished from T. squamatum by coarser areolation (2 – 3 areolae in 10 µm). In addition to these differences, Pantocsek distinguished between them by the proximity of internal costae to the pseudocelli-bearing polar elevations. In E. truanii and E. tschestnovii ‘ scales’ 1 come in close proximity to the pseudocelli, whereas in T. squamatum the costae are more distant from the pseudocelli. The polar elevations in E. truanii and E. tschestnovii were termed by Pantocsek (1889) as subcapitate and capitate, respectively. Our observations indicate that the areolation density and the extent of internal costae are highly variable, especially the number of areolae in 10 µm, which appears to be size-dependent. With respect to the valve face convexity in E. truanii and E. tschestnovii, the specimens Pantocsek examined may have been positioned with their valve faces pointing away from the observer, in which case it would be easy to mistake concavity for convexity. All specimens examined herein have deeply depressed valve faces. Unfortunately, no type slides for any of the three species in question are extant in BP (K. Buczkó, personal communication, 2013). These observations are partly corroborated by Brun (1896: 245), who suggested that T. squamatum and E. tschestnovii were conspecific and closely related to Triceratium decorum Greville (1862 a: 92) and T. pectinatum Greville (1862 a: 92). It is not possible to explore this idea, as neither T. decorum nor T. pectinatum have been examined in SEM, and Greville’s type specimen for T. pectinatum has not been found (Williams 1988: 57). Brun complicated matters by proposing the variety T. squatum var. radiata Brun (1896: 245). Brun’s diagnosis suggests that valves of his variety are smaller than those of the species and have radially arranged areolae. Both the specimens examined here, however, and Pantocsek’s illustrations show generally radial areolation patterns. Based on the range of size variation, we do not think the separation of T. squamatum var. radiata is warranted. Thus, all three taxa proposed by Pantocsek noted above are within the range of morphological variation for a single species. Since T. squamatum has priority over E. truanii and E. tschestnovii, the new combination Entogoniopsis squamata is proposed. Examination of Pantocsek’s type material from Kusnetzk may add to these conclusions.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0971FB15FF2FFB25FA762336.taxon	description	(SEM: Figs 120 – 124; LM: Figs 125 – 127)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0971FB15FF2FFB25FA762336.taxon	materials_examined	TYPE (here designated): — South Tasman Rise, Southwest Pacific Ocean, DSDP sample 29 - 281 - 16 - 3, 75 - 77 cm (SZCZ 21078, holotype! = Fig. 127). Valves triangular, with acute to slightly rounded poles, and straight sides (Figs 120, 125 – 127). Entire valve face is depressed (Fig. 122), apart from raised poles with prominent pseudocelli (Figs 120, 122). A hyaline marginal ridge extends between each pseudocellus. Whole valve face is perforated by radially oriented poroid areolae occluded by volae (Figs 120, 123, 125 – 127). Marginal zone with small areolae, more closely spaced than those within the central depression (Figs 125 – 127), which are also variable in size and shape. A well-defined ring of rimoportulae is located in the valve face centre or around the central area (Fig. 120), with internal openings in the form of slits across slightly raised papillae (Fig. 121). Mantle steeply downturned, shallow, perforated by volate poroid areolae arranged in rows parallel to the pervalvar axis (Fig. 124). Margin of the mantle smooth, hyaline, slightly expanded inwardly (Fig. 124). The expanded mantle margin on valve interior gives rise to a complex network of internal costae that intersect at multiple angles, forming a series of prominent ‘ loops’ along each margin of the valve face (Figs 120, 124). The costae reach a short distance toward the centre of the valve face (Fig. 120), and are manifested on the valve face exterior by defining sectors, in which occur the marginal areolae (Figs 125 – 127). Measurements (n = 8): average side length: 211.6 – 359.8 µm; 1 costa in 10 µm measured along the valve face margin; 2 areolae in 10 µm in marginal sectors; 1.5 – 2 areolae within the central depression; 9 – 18 rimoportulae; 4 – 5 porelli in 10 µm.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0971FB15FF2FFB25FA762336.taxon	etymology	Etymology: — Named for its superficial resemblance to E. squamata and the presence of the prominent central ring of rimoportulae. Geographic and stratigraphic distribution (Fig. 10, sites 12, 19): Middle Eocene: Falkland Plateau, South Atlantic: Vema Cruise 17, Core 107: BM stub P. 1274 (Figs 120 – 124). 1. Pantocsek referred to the scaly pattern produced by the internal costae, hence the Latin epithet ‘ squamatum’ 20 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. Late Eocene: South Tasman Rise, Southwestern Pacific Ocean: DSDP Site 281, Cores 281 - 14 through 281 - 16: SZCZ 15073, 21078 (Figs 125 – 127), 22054 – 22057. Observations: — Despite numerous hours of careful search, only a single specimen revealing the structure of the valve interior was available for SEM examination. Therefore, external openings of the rimoportulae were not observed in SEM, but LM observations strongly suggest that the external openings are in the form of simple slits.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0977FB16FF2FFCA2FF122388.taxon	description	(SEM: Figs 149 – 154; LM: Figs 155 – 160)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0977FB16FF2FFCA2FF122388.taxon	materials_examined	TYPE: — ‘ Moron deposit, Province of Seville’. Triceratium stokesianum var. moravica Grunow in A. W. F. Schmidt (1886 b: taf. 112, fig. 19). TYPE: — ‘ Brünn’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0977FB16FF2FFCA2FF122388.taxon	description	Geographic and stratigraphic distribution (Fig. 10, sites 18 *, 22 – 25, 28, 33 *; questionable records indicated with asterisks): (a) specimens: Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: BM coll. Adams GC 3419. 2 Middle Miocene: Szakal, Hungary: BM 63751 (Figs 156 – 157), BM coll. Adams C 921; Szurdokpüspöki (‘ Castel’), Hungary: BM stub P. 1352 (Figs 150 – 152), BM coll. Adams 741 (Fig. 159); Pôtor, Slovakia (formerly Szent-Peter, Hungary): BM stub E. 9 (Figs 149, 153 – 154), BM coll. Adams F 1387. Middle-late Miocene: Morón de la Frontera, Spain: BM coll. Adams TS 253. 2. Questionable: labelled as T. stokesianum but is probably Entogoniopsis pseudonervata comb. nov. (see below). 22 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. Late Miocene: Sendai, Japan: BM coll. Adams GC 3422. Age unspecified, presumably Miocene: Hungary: BM 63749 (Figs 155, 160), BM 63750 (Fig. 158). (b) records: Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Grove & Sturt (1887 c: 137), De Lautour (1888: 310), Tempère & Peragallo (1915: 396). These records are questionable, as there are no other records of Entogoniopsis stokesiana from sediments older than the Miocene time interval. As E. stokesiana is easily confused with E. pseudonervata (see below), we suggest that the reports by Grove & Sturt (1887 c), De Lautour (1888) and Tempère & Peragallo (1915) represent records of E. pseudonervata. Early Miocene: Horné Strháre (formerly Felsö-Esztergály), Slovakia: Pantocsek (1886: 54); Modrý Kameň, Slovakia (formerly Kékkö, Hungary): Pantocsek (1886: 54), Tempère & Peragallo (1915: 149); Marmorito, Italy: Tempère & Peragallo (1915: 197). Late early Miocene: Mogyoród, Hungary: Pantocsek (1886: 54). Early-middle Miocene: Brno region, Moravia, Czech Republic (formerly Brünn, Mähren): Cleve (1885: 169, pl. 12, figs 5 a, b), Schmidt (1886 b: taf. 112, fig. 19), Hustedt (1959 a: taf. 472, fig. 5); Přibice, Moravia, Czech Republic (formerly Pribitz, Mähren): Hustedt (1959 a: taf. 472, figs 4, 6).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0977FB16FF2FFCA2FF122388.taxon	description	Late Miocene: Sendai, Japan: Schmidt (1890: taf. 159, fig. 2), Tempère & Peragallo (1915: 15). Recent: Mann (1925: 46) recorded this species as extant in the Philippine Islands, transferring it to Biddulphia in the process. We consider Mann’s report as questionable, as there are no other records of E. stokesiana from sediments younger than the Miocene time interval. Similarly, we observed a specimen on slide BM coll. Adams GC 3421, labelled as coming from Manila, Philippine Islands. This diatom has a large tripolar valve with high polar elevations and short, evenly spaced internal costae located along each margin. The central area of the valve face is conspicuously domed, hence it does not appear to be E. stokesiana. This supports the notion that the occurrences of E. stokesiana are restricted to the Miocene. Observations: — Based on material from Morón, Spain, Greville (1866 a: 8) noted that in his specimens of Triceratium stokesianum, the ‘ veins’ protruding from the two adjacent sides of the valve are connected. Further, Greville suggested some morphological variation in the number of costae, and in the distance that they reach toward the valve face centre. From the Brno region of Moravia, Czech Republic, Cleve (1885: 169) reported specimens with internal costae that do not fuse. The description in Cleve (1885) emphasizes the variability in the pattern formed by internal costae. Grunow (1887 a: 66), however, suggested that the specimen reported by Cleve (1885) differs from that of Greville (1866 a), such that the former should be treated as new variety: T. stokesianum var. moravica Grunow (in Schmidt 1886 b: taf. 112, fig. 19). Grunow (1887 a; see also Schmidt 1886 b, taf. 112, fig. 19) did not specify the morphological features that distinguish this variety from the species, but presumably he meant the variations in the number and length of the internal costae. Hustedt (1959 a: taf. 472) found these taxonomic concepts confusing and proposed the following distinction: (1) T. stokesianum has undulate valves, with one raised sector per side bound by a pair of divergent internal costae; (2) T. stokesianum var. moravica has a circumferential depression around a raised valve face centre and the internal costae are generally convergent. In addition to valve view figures that support this distinction (taf. 472: figs 1 – 2, 5 – 6), Hustedt figured both in oblique or girdle view (1959 a: taf. 472, figs 3 – 4). The oblique view of his T. stokesianum shows a clear trifolium in the centre (taf. 472, fig. 3); the girdle view of T. stokesianum var. moravica shows a raised area in the middle of the side facing the observer, which also suggests a presence of a trifolium. Therefore, Hustedt (1959 a) may have mislabelled his girdle view figure of T. stokesianum var. moravica. Our examination of specimens from Morón, Sendai, and a number of Central European localities shows that the arrangement of internal costae in E. stokesiana is highly variable. A single specimen from Morón examined in this study (BM coll. Adams TS 253) shows faint connections between pairs of internal costae arising from adjacent valve margins. This specimen is also convex-sided, which is another character that agrees well with Greville’s description. However, SEM examination of the valve interior in a specimen from Szurdokpüspöki (‘ Castel’) (Fig. 150) shows that such connections are found also in specimens from other localities, and that they are merely local thickenings of the valve face. Therefore, whether the internal costae appear to fuse or not depends primarily on toward which side these costae are deflected. Specimen in Fig. 156 shows the presence of such thickened areas ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 23 between a pair of costae adjacent to one pole, and between pairs of neighbouring costae. Therefore, given the high degree of morphological variation in E. stokesiana, we see no use in defining infraspecific taxa based on the pattern formed by the internal costae alone. None of the specimens examined in this study showed a circumferential depression comparable to that reported by Hustedt (1959 a). As in the case of Trilamina wittiana (see below), we suggest that the morphological variation in E. stokesiana may be further complicated if this species formed heterovalvar frustules. Until intact frustules have been examined, we propose to synonymize the species and the variety.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	description	nov. (SEM: Figs 193 – 198, LM: Figs 199 – 204)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	materials_examined	TYPE: — ‘ Honduras’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, Cambridge Estate’ (BM 2972, holotype!). Triceratium grave A. W. F. Schmidt (1882: taf. 77, fig. 17). TYPE: — ‘ Jérémie’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	materials_examined	TYPE: — ‘ Maria Madre Island, Tres Marias Group, Mexico’ (CAS 200035, holotype!).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	description	Geographic and stratigraphic distribution (Fig. 10, sites 14, 21, 26, 29 – 37, 39 *; questionable records indicated with an asterisk): (a) specimens: Middle Eocene-early Miocene: Barbadoes: BM coll. Adams: H 907, TS 756. Oceanic Formation outcrops at Cambridge Estate: BM 2972, BM coll. Adams F 1389; Conset: BM stub P. 1338 (Fig. 197); Joe’s River: BM stub P. 1351 (Figs 193, 195), BM coll. Adams TS 927; Newcastle: BM 65915; Springfield: BM coll. Adams: Bess 343, GC 3431. Early Miocene: Jeremié, Haiti: BM coll. Adams: J 661, TS 543 (Fig. 202). Late Miocene: Maria Madre Island, Mexico: BM stub P. 1335 (Figs 194, 196, 198), BM coll. Adams TS 582 (Fig. 204). Recent: Campeche Bay: BM 55099; Colón, Panama: BM 63761 (Fig. 200), BM coll. Adams: F 1390, F 1391; Manila, Philippine Islands: BM 63765 (Figs 199, 201); Philippine Islands: BM coll. Adams TS 507 (label reads: ex Mann); Toamasina / Tamatave, Madagascar: BM coll. Adams TS 451 (Fig. 203). (b) records: Early Miocene: Jeremié, Haiti: Schmidt (1882: pl. 77, fig. 17), Truan & Witt (1888: 22, pl. VII, figs 13, 19, 21).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	description	Recent: Campeche Bay: Cleve (1878: 17); Honduras (presumably from mahogany logs): Brightwell (1856: 275, pl. XVII, fig. 15); Philippine Islands: Mann (1925: 46); Galapagos Islands and Virgin Islands: Cleve (1878: 17); off northern Mauritius: Desikachary (1989: 12, pl. 805, figs 1 – 6). Age unspecified, presumably Recent: Dredgings in the area of Galapagos Islands and in the Bering Sea: Mann (1907: 296). 3 Observations: — Entogoniopsis tabellaria has a high degree of variation in the number of internal costae per side, which is dependent on the valve diameter; the polar elevations vary from flat to convex. It is also unusual in having 3. The Bering Sea record is questionable: all verified records of E. tabellaria are from tropical and subtropical latitudes. No figures were provided by Mann (1907), and therefore verification is not possible. ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 27 distinctly domed vela. Apart from E. tabellaria and E. inflata, all other species of Entogoniopsis documented so far have flat, slightly sunken cribrate or volate occlusions to the areolae.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0978FB1AFF2FFF17FDF12497.taxon	description	From early Eocene sediments of the middle Urals area, Schibkova (in Krotov & Schibkova 1959: 120) proposed T. tabellarium var. areolata. No specimens of this variety were available to us; the description makes no mention of valve face undulations that could indicate the presence or absence of the central trifolium. In comparison to T. tabellarium var. diplosticta, Schibkova states that T. tabellarium var. areolata lacks ‘ lines’ along the margin. The figure provided by Schibkova (Krotov & Schibkova 1959: fig. 3.3), however, appears to show faint, short transverse lines along each margin of the valve face, which may represent internal costae. Given these uncertainties, the relationships of T. tabellarium var. areolata cannot be clarified until the material of Krotov & Schibkova (1959) has been reexamined. No specimens of extant E. tabellaria were available for SEM examination. However, the morphology of the specimens from Campeche, Philippines, Panama and Madagascar examined in LM is consistent with those of Miocene age, e. g., from Madria Madre Island, Mexico and from Java. Valves of the same general morphology are also found in the Oceanic Formation sediments exposed in Barbadoes (Figs 193, 195, 197); the degree of pseudocellus convexity (Figs 195 – 196) is the only morphological feature that distinguishes specimens from Barbadoes from those from the Miocene deposits. This, in our opinion, does not warrant separation and therefore we interpret them as conspecific.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFA37FF17251E.taxon	materials_examined	TYPE: — ‘ Barbadoes earth’.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097FFB1CFF2FFA37FF17251E.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit’. We interpret Triceratium truncatum and Triceratium productum to fall within the range of variation of the same species.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096FFB0FFF2FFAE2FC36251D.taxon	description	(SEM: Figs 52 – 57; LM: Figs 58 – 68)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096FFB0FFF2FFAE2FC36251D.taxon	materials_examined	TYPE: — “ Barbadoes earth ”. Biddulphia venosa (Brightwell) Grunow in Van Heurck (1883: pl. CXIII, fig. 3). Triceratium venosum f. parva A. W. F. Schmidt (1886 a: taf. 94, fig. 12). TYPE: — “ Barbadoes, New Castle ”. Triceratium venosum var. major Laporte & Lefébure (1929: pl. X, fig. 71). TYPE: — “ Barbades. Rare ”.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096FFB0FFF2FFAE2FC36251D.taxon	description	14 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. parallel to the pervalvar axis on the mantle (Figs 52 – 53, 56). A network of robust costae is located on the valve interior (Fig. 53). Usually, three prominent, long costae radiate from the central rimoportula toward the polar elevations (Figs 53, 55), and numerous shorter, transverse costae branch from the longest ones at more or less regular intervals (Figs 53, 57). The transverse costae extend toward the valve margin and continue down the mantle (Fig. 57). Internal costae on the mantle are associated with short, inconspicuous external costae located on the mantle part adjacent to the valve face (Fig. 56). Measurements (n = 40): average side length: 46.5 – 139.1 µm; 3 – 4 areolae in 10 µm; 2 – 4 costae in 10 µm measured along the valve face margin; pseudocelli: 8 – 10 porelli in 10 µm. Geographic and stratigraphic distribution (Fig. 10, sites 2, 3 *, 9, 14, 18; questionable records indicated with an asterisk): (a) specimens: Late Palaeocene: Kusnetzk, Russia: BM coll. Adams M 89. Middle Eocene: Western North Atlantic Ocean: ODP Site 1050 A, Core 1050 A- 2 H (Fig. 68); ODP Site 1051: SZCZ 16102 A, 16105 A, 17090 D, 17094 B, 17096 C, 17099 C, 17100 A, 17105 A, 17110 C, 17113 D, 17119 C, 17122 D, 17942 B, 17943 B, 17944 B, 17946 D, 17951 D, 17991 A, 18003 B, 18234 B, 18236 B, 18238 C, 18242 C, 18248 B, 18263 A, 18704 B, 18836 A, 18837 A, 18839 B, 18846 D, 18850 D, 18861 C, 18891 A, 18893 C, 18895 D, 18898 C, 18904 C, 18905 C, 18906 D. Middle Eocene-early Miocene: Barbadoes: BM 35040 (Fig. 59), BM 68793 (Figs 60, 66), BM coll. Adams TS 266; Oceanic Formation outcrops at: Mt Hillaby: BM coll. Adams TS 491; Joe’s River: BM stub P. 1281 (Figs 52, 54, 56), BM 35041, BM 35926, BM 63792 (Fig. 58), BM 65924, BM coll. Adams TS 856 (Fig. 62); Springfield: BM 38101, BM 38104 – 38108 (Fig. 67); Newcastle: BM 63791 (Fig. 61); Chalky Mount: BM 38103. Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: BM stub P. 1274 (Figs 53, 55, 57), BM 11107 (Fig. 63), BM 68793 (Fig. 64), BM coll. Adams: G 11, TS 315 (Fig. 65); Oamaru Diatomite outcrop at Jackson’s Paddock: BM 63794. (b) records:	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E096FFB0FFF2FFAE2FC36251D.taxon	description	Late Eocene-earliest Oligocene: Oamaru, Otago, New Zealand: Crosby & Wood (1958: 508); Oamaru Diatomite outcrops at: Cormack’s Siding: Grove & Sturt (1886: 327), De Lautour (1888: 311), Desikachary & Sreelatha (1989: 270); Bain’s Middle and Division Hill: Desikachary & Sreelatha (1989: 270); Flume Gully: Doig (1991: 125). Observations: — The most variable morphological feature in E. venosa is the valve outline. Valves examined here range from those with broadly rounded poles (Fig. 66) to those with produced, even subcapitate poles (Figs 60, 62). Variation in valve outline appears to be independent of valve diameter. Considerable variation is also noted in size of the areolae on the valve face. The general pattern of valve face areolation in E. venosa is consistent, but numerous valves examined here in LM showed minor differences. Some rare specimens appear to lack the central rimoportula (Fig. 64), but generally both the number and location of rimoportulae are consistent morphological features of E. venosa. Additionally, some specimens of E. venosa have an expanded internal hyaline central area associated with the central rimoportula (Figs 63 – 66), similar to the central area seen in E. polycistinora (Figs 39, 43). In the latter, the central area hosts multiple rimoportulae, whereas E. venosa consistently has a single process; therefore, the function of the internal plate in E. venosa is unknown, but its presence suggests a close phylogenetic relationship between these two species. Close examination of the internal costae in E. venosa shows that the prominent radial costae extending between the central rimoportula and the poles are in fact composed of segments connecting the transverse costae. In some of the specimens examined, some segments are missing (e. g., Fig. 65), suggesting that the transverse costae were formed earlier in valve morphogenesis than the radial costae.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0979FB1DFF2FF946FE8521AC.taxon	description	Frustule subrectangular in girdle view (Fig. 209), valve outline tripolar (Figs 205, 207), with acute poles and straight sides. Valve face undulate, with a distinct trifolium bearing a small hyaline area at the centre (Fig. 205), within which there may or may not be rimoportulae (Figs 205, 207). Low elevation that bears a prominent, hooked linking spine at its distal face is present at each pole (Figs 205, 210 – 211). A hyaline marginal ridge extends between each polar elevation (Fig. 205). In the two sibling valves the hyaline marginal ridge is level with the summits of the polar elevations (Fig. 209), and there are additional small interlocking linking devices in even intervals along the crest of the ridge (Figs 209 – 210). The valve that lacks a sibling has a lower marginal ridge that appears to follow the valve face undulations, and without the linking devices 28 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. (Fig. 205). Radially oriented, locally curved hyaline costae are present on the valve face exterior (Fig. 205). Whole valve face is perforated by poroid areolae with radial, slightly domed cribra (Fig. 211). Mantle is shallow, steeply downturned, perforated by rows of poroid areolae parallel to the pervalvar axis (Fig. 207), slightly divergent closer to the poles. Margin of the mantle is hyaline, smooth, and slightly expanded inwardly (Fig. 207). Along each side on the valve interior, there is a series of robust costae that arise from the expanded mantle margin and project a short distance toward the valve face centre (Fig. 207). At least two copulae were observed. Valvocopula is closed, relatively deep, with only two poorly defined rows of poroid areolae near the advalvar margin (Figs 205, 209). Copula is closed, shallower than the valvocopula, mostly hyaline, with only a single row of poroids near the advalvar edge (Figs 205, 209). Remnants of a third girdle element can be seen projecting from underneath the copula, but are not interpretable (Fig. 211). Geographic and stratigraphic distribution (Fig. 10, site 23): Middle Miocene: Szurdokpüspöki (‘ Castel’), Hungary: BM stub P. 1352 (Figs 205 – 211). Observations: — Despite considerable effort, only a single specimen was found and that could only be examined in SEM. The presence of linking spines and secondary interlocking linking devices located along the crest of the hyaline marginal ridge, heterovalvy with respect to the presence or absence of rimoportulae, the absence of pseudocelli, and the lack of compelling evidence for valvocopula attachment to the internal costae suggest that this taxon might represent a separate genus. However, given the insufficient data, and the lack of further specimens with which to typify it, we are reluctant to describe this as new species and provisionally identify it as an unknown species of Entogoniopsis. New combinations	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0943FB20FF2FFC19FC772634.taxon	materials_examined	TYPE: ‘ Barbadoes deposit, from Cambridge estate’ (BM 3281, holotype!).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0943FB20FF2FFBDCFBE726F7.taxon	materials_examined	TYPE: ‘ Cambridge, Barbadoes’ (BM 2757, holotype; specimen lost, Williams 1988: 55).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097CFB1EFF2FFC65FC06212B.taxon	description	(SEM: Figs 217 – 222; LM: Figs 223 – 229)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097CFB1EFF2FFC65FC06212B.taxon	materials_examined	TYPE: — ‘ Barbadoes deposit, Cambridge estate’ (BM 3198, holotype! = Fig. 228; Williams 1988: 56). Frustules rectangular to subrectangular in girdle view, valves tripolar (Fig. 223 – 229) with deeply concave sides and wedge-shaped (Figs 227, 229) to linear projections (Figs 223, 228), rounded or tapering near the pole. An inconspicuous pseudocellus is located at each pole (Fig. 219). The valve centre, with a triangular or hexagonal outline (Figs 224 versus 223), is convex and mostly hyaline, with few irregular rings of sparse poroid areolae near the margins (Fig. 218). The marginal parts of the projections are perforate, with poroid areolae arranged in transverse, short, parallel rows that continue down the mantle (Figs 218 – 219). Size of areolae diminishes toward the mantle margin (Figs 218 – 219). Valve face-mantle transition is gradual, mantle is steep and shallow (Figs 217 – 218). The central part of each projection bears a prominent, depressed hyaline area tapering toward the pole (Figs 217 – 218). A prominent network of robust costae (Figs 220 – 221) is present on the valve interior. The most prominent of these costae are associated with the depressed hyaline areas within each projection (Fig. 220). Tapering costae give rise to transverse, shorter costae that continue down the mantle and terminate at the inwardly expanded hyaline mantle margin (Fig. 221). On the valve interior, each side of the central area is associated with a robust costa or a transverse fold (Figs 220 – 221). A single, inconspicuous rimoportula is placed next to one of the folds (Fig. 220). On the interior, it is a slit between two slightly raised lips (Fig. 220). The rimoportula lacks a tube, and thus it is indistinguishable from other perforations on the exterior (Fig. 218). Valvocopula is closed, comparable in depth to the mantle, attached to the valve by means of a fossa that embraces the inwardly expanded hyaline margin of the mantle, large clasping devices attached at mid-point of each side to internal costae or folds that line the central area (Fig. 221), and small clasping devices located in regular intervals along the sides of each projection (Fig. 222). The small clasping devices attach to each of the transverse costae that run down the mantle (Fig. 222). Valvocopula is mostly hyaline, with a single row of poroids located next to the advalvar edge (Fig. 221), interrupted at mid-point of each side, due to the presence of the large clasping devices. Measurements (n = 13): average side length: 44.7 – 124.5 µm; 3 – 4 costae in 10 µm measured along the valve face margin; 3 – 5 areolae in 5 µm measured across a projection; 6 – 7 rows of areolae in 10 µm, measured along a projection. Geographic and stratigraphic distribution (Fig. 10, sites 14, 18): (a) specimens: Middle Eocene-early Miocene: Oceanic Formation outcrops at Cambridge Estate: BM 3198 (Fig. 228, holotype); Joe’s River: BM 63659 (Fig. 223), BM 65907, BM coll. Adams H 772. ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 31 Late Eocene-earliest Oligocene: Oamaru, New Zealand: BM stub P. 137 (Figs 217 – 222), BM coll. Adams TS 694 (Fig. 224), BM 63658 (Fig. 225), BM 61163, BM 61164 (Figs 226 and 229), BM coll. Adams GC 3331, GC: 3332 (Fig. 227), H 302, J 3666. (b) records:	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097CFB1EFF2FFC65FC06212B.taxon	description	Observations: — Based on superficial resemblances, Greville (1865 a: 8) suggested that T. nitescens was ‘ allied’ to Triceratium lobatum Greville (1863: 233). Prior to publishing the description of T. nitescens, Greville proposed a number of other similar taxa in addition to T. lobatum: Triceratium westianum (Greville 1861 a: 43, pl. IV, fig. 11), Triceratium rotundatum (Greville 1861 c: 75, pl. IX, fig. 6), Triceratium denticulatum (Greville 1863: 233, pl. IX, fig. 14) and Triceratium obesum (Greville 1864 b: 90, pl. XIII, fig. 11). None of these, however, were indicated by Greville to be a possible close relative of T. nitescens. Of all these taxa, T. westianum has been available for SEM examination (see below). However, the presence of extensive hyaline areas and an overall similar pattern of internal costae in all of these species has prompted their transfer to Trilamina.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0943FB20FF2FFAA4FD752402.taxon	materials_examined	TYPE: ‘ Barbadoes deposit, from Cambridge estate’ (BM 3078, holotype! = Fig. 252).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097CFB1FFF2FFF17FDF820CF.taxon	materials_examined	TYPE: — Trilamina nitescens (Greville) J. Witkowski, P. A. Sims, Strelnikova & D. M. Williams, comb. nov. [= Triceratium nitescens Greville (1865 a: 8, pl. II, fig. 19)]. Frustules rectangular in girdle view, valves tri-to quadripolar, with deeply concave sides and tapering or broadly lobed projections distinctly cut off from the mostly hyaline valve centre. Rimoportulae, if present, indistinct and located offcentre within the central area. Central part of each projection depressed, either hyaline or perforated by poroid areolae. Marginal part of the valve raised, and perforated by poroid areolae arranged in short rows parallel to pervalvar axis. Indistinct pseudocelli located at the poles. Valve face-mantle junction gradual, mantle steep, shallow. Internal costae present along the sides of projections, sometimes also associated with the depressed areas within projections; additional internal costae occur in some species on the underside of the central area-Valvocopulae are closed, comparable in depth to the mantle, mostly hyaline, with few rows of poroids next to the advalvar edge. The valvocopula attachment mechanism consists of (1) small clasping devices that attach to the internal costae; (2) fossa that attaches to the inward expansion of the hyaline mantle margin; and (3) large clasping devices that attach to the internal folds associated with the margins of the central area (a synapomorphy for this genus). Stratigraphic range: middle Eocene-middle Miocene Habitat: Neritic, likely benthic.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0943FB20FF2FF9AEFCF024EE.taxon	materials_examined	TYPE: ‘ Barbadoes deposit’ (BM 2111, holotype! = Fig. 251).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0942FB20FF2FFD44FC062172.taxon	description	(SEM: Figs 239 – 242; LM: Figs 243 – 248)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0942FB20FF2FFD44FC062172.taxon	materials_examined	TYPE: ‘ Barbadoes deposit’ (BM 2029, holotype! = Fig. 245; Williams 1988: 60). Valves tri-to quadripolar (Figs 243 – 248), with broadly rounded projections, usually distinctly capitate (Figs 244 – 247), and deeply concave sides. Valve face undulate: central area and the marginal zone are raised, whereas the central part of each projection is depressed (Fig. 240). The central area bears a small hyaline field, with a single rimoportula offset toward one of the projections (Figs 239, 242). In the marginal part of the central area there are few poorly defined rings of areolae (Figs 243 – 248). The projections are perforated by poroid areolae: those within the depressed part are slightly larger, while the marginal areolae are slightly smaller. A field of smaller areolae that possibly represents a pseudocellus is located at each pole (Fig. 241). The larger areolae are arranged in rows parallel to the major axis of the projection. The marginal areolae are aligned in rows parallel to the pervalvar axis, and continue down the shallow mantle (Fig. 240). Valve face-mantle transition is gradual; a thickened, hyaline area comparable in width to the diameter of the central area is located at the mid-point of each side (Fig. 240). Mantle margin is smooth, hyaline, slightly expanded inwardly (Fig. 240). On the valve interior, there are numerous costae radiating from the distal part of the depressed area within each projection (Figs 243 – 248). Valvocopula is closed, its depth comparable to that of the mantle (Fig. 240). Large clasping devices on the valvocopula attach to the thickenings at the mid point of each side, and the fossa attaches to the inwardly expanded margin of the mantle (Fig. 240). Except for a single row of poroid areolae, interrupted at the locations of the large clasping devices and located adjacent to the fossa, entire valvocopula is hyaline (Fig. 240). Measurements (n = 8): average side length: 32.2 – 60.3 µm; 3 – 4 costae in 10 µm; 3 – 3.5 areolae in 10 µm. Geographic and stratigraphic distribution (Fig. 10, sites 9 *, 14, 15 *, 16; possible questionable records are indicated with an asterisk): (a) specimens: Eocene: Mascarene Ridge, Indian Ocean, dredging DODO- 123 - D 1: BM stub P. 1324 (Figs 239 – 242). Middle Eocene-early Miocene: Barbadoes: BM 2029 (Fig. 245, holotype), BM 52820 (Fig. 243), BM coll. Adams F 1414 (Fig. 246). Oceanic Formation outcrops at Springfield: BM 38130, BM 63799, BM coll. Adams GC 3466 (Fig. 244), J 4252 (Fig. 247); Newcastle: BM coll. Adams F 1413 (Fig. 248). (b) records: Middle Eocene: Northern Indian Ocean: ODP Site 707, Core 707 C- 9 R: Fenner & Mikkelsen (1990: 448, pl. 3, fig. 5) and ODP Site 713, Core 713 A- 9 R: Fenner & Mikkelsen (1990: 448, pl. 3, fig. 6). 4	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0942FB20FF2FFD44FC062172.taxon	description	4. These records are questionable and judging by valve outline, they likely represent Trilamina tripes (Site 707) and Trilamina figurata (Site 713). ENTOGONIOPSIS GEN. NOV. AND TRILAMINA GEN. NOV. (BACILLARIOPHYTA) Phytotaxa 209 (1) © 2015 Magnolia Press • 33 Observations: — The poor preservation of the specimen examined here in SEM does not allow inspection of the radial internal costae for the attachment mode of its valvocopula. However, small clasping devices are present in both T. nitescens and T. wittiana, with which T. westiana shares a number of morphological features. Therefore, we assume that this species also has a three-fold mechanism for valvocopula attachment. Regardless of the locality, valves of T. westiana are consistently small, with an average side length less than 35 µm. Although no specimens of Trilamina tripes (Fig. 249) were available for SEM examination, its valve morphology resembles T. westiana. The sparse records of T. tripes show that its valve diameter is close to the upper limit of the variation range in T. westiana. Both species have areolated depressed parts of the projections, and a small hyaline field within the central area. It is unclear, however, whether rimoportulae are present in T. tripes. Given the uncertainty in dating of the deposits from which both T. westiana and T. tripes are reported, it is impossible to state with confidence whether the stratigraphic ranges of these species overlap. However, T. tripes could potentially belong within the range of variation of T. westiana (but see also discussion under T. wittiana). New combinations Trilamina denticulata (Greville) J. Witkowski, P. A. Sims, N. I. Strelnikova & D. M. Williams, comb. nov. (LM: Fig. 250)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E0942FB20FF2FFD44FC062172.taxon	materials_examined	TYPE: ‘ Barbadoes deposit, from Cambridge estate’ (BM 2763, holotype! = Fig. 250).	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097DFB21FF2FFCDCFD2F2096.taxon	description	comb. nov. (SEM: Figs 230 – 235; LM: Figs 236 – 238)	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097DFB21FF2FFCDCFD2F2096.taxon	materials_examined	TYPE: — ‘ Jérémie’. Valves tripolar, with tapering or broadly rounded projections, and deeply concave sides (Figs 236 – 238). Valve face undulate, with margins and the central area raised to equal level, and a prominent depressed area within each projection (Fig. 230). The central area and the depressed areas are hyaline (Figs 230 – 231), except for a poorly defined ring of sparse poroid areolae in the marginal part of the central area (Fig. 230), and few poroids next to the mid-point of each side (Figs 236 – 238). The raised valve face margin, and most of the mantle are finely areolated, with poroid areolae aligned in rows parallel to the pervalvar axis (Figs 230, 232). Areolae become slightly reduced at each pole, which may represent a poorly defined pseudocellus (Fig. 232). Mantle is shallow, steep, but with a gradual valve face-mantle transition (Figs 230, 232). At the mid-point of each side, a narrow hyaline area is present on the mantle (Fig. 230). Mantle margin is smooth, hyaline, slightly expanded inwardly (Fig. 233). On the valve interior, there is a series of robust costae radiating in regular intervals from each of the depressed hyaline areas (Figs 231, 234, 236 – 238). Additional, shorter costae are found on the underside of the raised central area (Fig. 231). Next to the junction of the central area and the projections, two large costae protrude from each of the depressed hyaline areas toward the mid point of the adjacent sides (Figs 231, 235). These large costae provide attachment for the large clasping devices located on the valvocopula (Fig. 235). Small clasping devices on the valvocopula attach to the remaining marginal costae (Figs 231, 233 – 234), and a fossa attaches to the inward expansion of the mantle margin (Figs 231, 233). Valvocopula is closed, its depth comparable to that of the mantle (Figs 230, 232, 234). A single row of evenly spaced poroid areolae, interrupted only where the large clasping devices are located, is immediately adjacent to the fossa (Fig. 234). Otherwise, the valvocopula is hyaline (Figs 230, 232 – 234). Measurements (n = 5): average side length: 49.8 – 87.0 µm; 3 – 4 costae in 10 µm measured along the valve face margin; 3 – 4.5 areolae in 5 µm measured across a projection; 8 – 10 rows of areolae in 10 µm, measured along a projection. Geographic and stratigraphic distribution (Fig. 10, sites 21, 26): (a) specimens: Early Miocene: Jérémie, Haiti: BM 63801 (Fig. 238). Middle Miocene: Wonosari-Series, Java, Indonesia: BM stubs P. 683 (Fig. 234), P. 1319 (Figs 232, 235), and P. 1333 (Figs 230 – 231, 233), BM 36216 (Fig. 236; label reads „ Reinhold sample AY- 12 ”), BM 63802 (Fig. 237). (b) records: Early Miocene: Jérémie, Haiti: Truan & Witt (1888: 23, pl. VII, fig. 16);.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
ED69878E097DFB21FF2FFCDCFD2F2096.taxon	description	32 • Phytotaxa 209 (1) © 2015 Magnolia Press WITKOWSKI ET AL. have been available for SEM examination. As figured in Truan & Witt (1888: pl. VII, fig. 16), specimens from the type locality have broadly circular projections, whereas valves from the Javan material (Reinhold 1937) consistently have tapering, wedge-shaped projections. A specimen on slide BM 63801 (Jérémie; Fig. 238) has a single row of poroid areolae located along the margin of the depressed hyaline area in each of the projections. No such perforations were observed in the specimens from Java. However, we see no additional morphological differences between the specimens from both localities. In particular, the morphology of the central area is identical in all examined specimens. Finally, the comparable age of both deposits, and their location in tropical latitudes, strongly suggests that specimens from Jérémie and Java are conspecific. The valve outline of T. wittiana from Jérémie is similar to that of Trilamina tripes comb. nov. (Fig. 249). The difference between the two is that the depressed parts of the projections in T. wittiana are hyaline or bear a single marginal row of poroids, whereas in T. tripes the entire central part of each projection is perforated. Given the large uncertainty in the age assignment of the various Barbadean deposits, it is impossible to confidently state that T. tripes (originally proposed from Mount Hillaby, Brun 1896: 246) is age-equivalent to T. wittiana. However, the valve outline similarity suggests that these two taxa may represent valves of heterovalvar frustules. Examination of intact frustules is required to verify this hypothesis.	en	Witkowski, Jakub, Sims, Patricia A., Strelnikova, Nina I., Williams, David M. (2015): Entogoniopsis gen. nov. and Trilamina gen. nov. (Bacillariophyta): a survey of multipolar pseudocellate diatoms with internal costae, including comments on the genus Sheshukovia Gleser. Phytotaxa 209 (1): 448-450, DOI: 10.11646/phytotaxa.209.1.1, URL: http://dx.doi.org/10.11646/phytotaxa.209.1.1
