identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
A5388564FFFF50205F2425FEFE65FE05.text	A5388564FFFF50205F2425FEFE65FE05.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Psammina yokosukae Gooday & Ishitani & Chen & Holzmann & Richirt & Seike & Yamashita & Tsuchiya & Nomaki 2025	<div><p>Psammina yokosukae sp. nov.</p><p>urn:lsid:zoobank.org:act: 8F06D3F4-DE75-46FD-8701-0D6C1379EB9E</p><p>Figs 2A–B, 3–4, 5D–G, 6–7, 8A–E; Supp. file 2: Figs S4A–B, S 5A–B, Supp. file 3: µCT video 1</p><p>Diagnosis</p><p>Species of Psammina with free, epifaunal test comprising system of undulating, sometimes fan-shaped, plates up to&gt; 5 cm in overall extent. In places, surface displays vague concentric zones defined by slightly raised lineations and traversed by low radial ridges. Micro-CT scans reveal internal structures, often rather poorly defined, corresponding to these surface features. Obvious apertures absent. Test wall friable, comprising mineral grains of varying sizes (generally &lt;100 µm) and occasional biogenic particles. Interior partly filled with agglutinated mineral grains that are larger (&gt;300 µm) than those constituting the wall. Stercomare forms branching and anastomosing strings and masses of variable width (~70 to&gt;200 µm) that occupy spaces between internal grains with no clear pattern. Granellare yellowish-brown, branching, thread-like (width ~10 to 34 µm), in places attached to grain surfaces; barite crystals rare.</p><p>Etymology</p><p>Named for the RV Yokosuka, support ship for the HOV Shinkai 6500.</p><p>Material examined</p><p>Holotype NW PACIFIC OCEAN – 32.5° N (west of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=143.77066&amp;materialsCitation.latitude=32.580166" title="Search Plazi for locations around (long 143.77066/lat 32.580166)">Kuroshio Extension Observatory</a> (KEO)) • 32°34.81′ N, 143°46.24′ E; depth 5509 m; 22 Oct. 2022; H. Nomaki leg.; Dive 1659 of HOV Shinkai 6500, core Red#3; GenBank accession nos: PP662675 to PP662677; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr691. The specimen is preserved in a core in 10% buffered formalin.</p><p>Paratype</p><p>NW PACIFIC OCEAN – 32.5° N (west of KEO) • 1 spec.; same data as for holotype; Dive 1659 of HOV Shinkai 6500, core Red#5; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr691. The specimen was frozen at sea and only available as fragments .</p><p>Description</p><p>Overall test morphology</p><p>The in situ seafloor photograph of the holotype (Fig. 2A) shows a group of about five main plate-like elements that are curved or undulating with sinuous and often upturned margins. The plates are orientated in different directions but merge together into one structure. Near one end of the test, there is an open-ended, chimney-like feature that is also visible in the shipboard photograph (Supp. file 2: Fig. S4A). The seafloor photograph of the paratype (TNS Pr693) is similar, although it comprises fewer, relatively larger plates (Fig. 2B).</p><p>The recovered holotype measures 5.5 cm × 3.7 cm in maximum extent when viewed from above (Fig. 3B) and extends ~ 2.7 cm above the sediment surface. The test is composed of plates that are ~0.60–1.0 mm (mean 0.81 ± 0.11 mm, n = 11) thick. It is strongly asymmetrical, with one side low lying, or even slightly buried under resedimented material (Figs 3A–B, 4C–D, G–H), and the other side upstanding. Supp. file 3: µCT video 1 gives a clear impression of its structure. The low-lying side comprises two mainly separate plates with strongly upturned rims. These, and particularly the larger of the two, merge with, and to some extent buttress, the more prominent upstanding part of the test. This part can be thought of as comprising two components. One is a branched plate, the upper part of which forms a U-shaped trough with undulating side (Fig. 3A, E). One side of the trough merges with the second component, a fairly flat semicircular fan that is a visually conspicuous feature in Figs 3C, 4A–D, and shown in detail in Fig. 3D–G. In side view, the fan appears to be linked with the abovementioned tubular chimney-like structure (Fig. 3A–C, G), but the µCT video (Supp. file 3) clearly shows that this feature develops at the end of one side of the U-shaped trough, after it has merged with the fan-shaped plate. Another approximately circular, somewhat larger but less regular feature is located in the central part of the test. It is largely obscured by other test elements but can be seen at a few points in the video.</p><p>Surface ornamentation, test structure and composition</p><p>In light photographs of the holotype, the test surface displays a vague pattern of concentric zones. These can be seen most clearly under very oblique lighting, notably on the prominent semicircular plate (Fig. 3G; Supp. file 2: Fig. S5B). Where the boundaries between the zones are most clearly developed (Fig. 3G), they appear to be shallow steps, suggesting that later zones overlap earlier ones. Superimposed on these zones are low radial ridges separated by shallow furrows, which are equally indistinct. The same pattern is clearer in μCT images of the holotype than in visible light. When the test is rendered more transparent in μCT scans, it is almost entirely occupied by concentric zones and radial features. Although the pattern of lineations is obvious on the semicircular plate (Fig. 3C–D), the individual features are often indistinct and discontinuous.</p><p>The test wall is fairly friable, 180–320 µm (mean 267 ±56 µm, n = 15) thick, and light greyish in colour when dry. It is continuous around the margin and devoid of obvious apertures. Seen through the wall of the core tube, the surface is peppered with a variable density of larger grains (Fig. 3F–G; Supp. file 2: Fig. S5B). Under a stereo microscope, the surface appears noticeably granular and comprises a rather heterogeneous mixture of mineral grains of various sizes (Fig. 5D, F). Some larger ones are dark and there is a sprinkling of whitish and orange grains, but most are pale or transparent.</p><p>In SEM images of the paratype, the surface is quite uneven at a scale of 100–200 µm (Fig. 6A–D). The main constituents are mineral grains amongst which are scattered some biogenic remains. mainly radiolarian fragments, but also diatom fragments. The outer wall is not sharply delimited from the test interior (Fig. 6G). In addition to empty space, the interior is occupied by a variety of particles (‘internal xenophyae’) of different shapes and sizes that are dominated visually (although not numerically) by grains that are larger (200–300 µm) than those forming the outer layer (Fig. 7A–D). These include sharp-edged shards of volcanic glass, and porous and fibrous particles, possibly also of volcanic origin, in addition to radiolarian fragments.</p><p>The relatively few test fragments in which the interior was examined directly did not display welldefined partitions corresponding to the internal structures visible in μCT renders (Fig. 3C–D). However, two or perhaps three poorly-defined internal partitions can be seen in an SEM image showing the interior exposed at the edge of a test fragment (Fig. 6A); a broken cross section of the test shows similar features (Fig. 6G). Vague radial structures are sometimes also discernible when the test interior is viewed by light microscopy (Fig. 5E). The poorly defined nature of these features probably reflects the fact that the internal agglutination is dominated bys coarse-grained particles. This probably also explains why individual radial features are sometimes indistinct and discontinuous in µCT renders (Fig. 3D).</p><p>Stercomare and granellare</p><p>In addition to internal particles, the test contains stercomare and granellare. The dark stercomare forms branches that to some extent anastomose, as well as more irregular masses (Figs 5G, 6F, 8A). At least in the test fragments examined, they did not follow any clear trend but occupied the irregular spaces between the large internal xenophyae. The branches vary greatly in width, from ~70 µm to more than 200 µm. They have a thin, transparent organic sheath, ~1 µm thick, that encloses masses of spherical to ovoid stercomata, each stercome ranging in size from 12 to 30 µm (typically 12–20 µm) (Figs 5G, 6H).</p><p>The granellare forms narrow thread-like strings, yellowish-brown in colour (a darker golden-brown when dried), that branch throughout the test interior (Figs 5G, 7A–D, 8A). They measure 10–40 µm in width (usually 14–25 µm; mean 20.7 ±6.17 µm, n=50) and occupy only a very small proportion of the interior space. The branches weave between the internal xenophyae and in places are attached to the surfaces of grains, from which they are difficult to dislodge. The organic sheath that surrounds the cytoplasm is ~0.1– 0.2 µm thick and shrinks creating longitudinal wrinkles when dried on an SEM stub (Fig. 7E). Granellare fragments viewed in glycerol under a compound microscope show the cytoplasm with small inclusions of various kinds, including a few refractive particles that are presumably mineral grains (Fig. 8D–E), although these do not resemble typical barite crystals. There was little evidence for intracellular barite crystals in SEM images. None were visible through the granellare walls (Fig. 7E). Various inclusions could be seen within the cytoplasm where the granellare tube was ruptured, but only a single crystal-like particle yielded EDS peaks for Ba (Fig. 7H), suggesting that barite crystals are not common.</p><p>Molecular characterisation</p><p>Psammina yokosukae sp. nov. branches as sister to Psammina tenuis Gooday &amp; Holzmann, 2020 . The grouping is supported by a BV of 89%. The sequenced fragment of the 18S gene of P. yokosukae contains 1038–1039 nucleotides and the GC content is 36 %.</p><p>Remarks</p><p>Phylogenetic reconstruction shows that Psammina yokosukae sp. nov. is closely related to P. tenuis . In terms of morphology, the granellare branches of both species are similar in being narrow ( P. yokosukae 12–22 µm; P. tenuis 10–30 µm) and attached for part of their length to internal test particles. The two species are also similar in having basically plate-like tests. An in situ photograph of the unique specimen of P. tenuis on the seafloor shows a relatively simple plate, strongly curved around a vertical axis, growing up vertically from the surface of a polymetallic nodule (Gooday et al. 2020a: fig. 5a therein). This is somewhat reminiscent of the holotype of P. yokosukae but is much simpler than the cluster of plates that make up the test of the new species. Concentric lines are more clearly developed in P. tenuis than in P. yokosukae, but there is no sign of the radial lineations that are dimly visible in the new species. These differences in test morphology support the genetic distinction between P. yokosukae sp. nov. and P. tenuis . Given the variability of many xenophyophores, an important caveat is that the overall test morphology is known in detail for only one specimen of both species. Nevertheless, the fact that the test wall is composed largely of radiolarian shells in P. tenuis but largely of mineral grains in P. yokosukae, and the presence of more numerous internal particles in P. yokosukae, supports the conclusion that these are different species.</p></div>	https://treatment.plazi.org/id/A5388564FFFF50205F2425FEFE65FE05	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gooday, Andrew J.;Ishitani, Yoshiyuki;Chen, Chong;Holzmann, Maria;Richirt, Julien;Seike, Koji;Yamashita, Momo;Tsuchiya, Masashi;Nomaki, Hidetaka	Gooday, Andrew J., Ishitani, Yoshiyuki, Chen, Chong, Holzmann, Maria, Richirt, Julien, Seike, Koji, Yamashita, Momo, Tsuchiya, Masashi, Nomaki, Hidetaka (2025): Integrative taxonomy of new xenophyophores (Rhizaria, Foraminifera) from the abyssal northwest Pacific. European Journal of Taxonomy 1004: 144-189, DOI: 10.5852/ejt.2025.1004.2973, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2973/13399
A5388564FFF5503C5F3322E7FDADFCAA.text	A5388564FFF5503C5F3322E7FDADFCAA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Psammina contorta Gooday & Ishitani & Chen & Holzmann & Richirt & Seike & Yamashita & Tsuchiya & Nomaki 2025	<div><p>Psammina contorta sp. nov.</p><p>urn:lsid:zoobank.org:act: E698FACF-6DE6-461F-95F3-EA971F1F8695</p><p>Figs 2C–D, 5A–C, 8F–G, 9–10, 22B; Supp. file 2: Figs S4C–D, S 5C–F; Supp. file 4: µCT video 2</p><p>Diagnosis</p><p>Species of Psammina with free, epifaunal test comprising complex, irregular system of variously shaped, often crookedly curved, plate-like elements up to ~ 60 mm in overall extent with no clear growth pattern or symmetry. Obvious apertures absent. Concentric zones sometimes present. Micro-CT scans reveal irregularly shaped, cell-like internal compartments. Test wall friable, comprising mixture of mineral grains of different sizes and colours. Stercomare forms branching and anastomosing strings of variable width that occupy internal spaces with no clear pattern. Granellare strands yellowish-brown, branching, thread-like (width ~14 to 40 µm), in places attached to grain surfaces; barite crystals not observed.</p><p>Etymology</p><p>From the Latin ‘ contortus ’, referring to the crooked and irregular appearance of different parts of the test.</p><p>Material examined</p><p>Holotype NW PACIFIC OCEAN – 32.5° N (west of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=143.77066&amp;materialsCitation.latitude=32.580166" title="Search Plazi for locations around (long 143.77066/lat 32.580166)">Kuroshio Extension Observatory</a> (KEO)) • 32°34.81′ N, 143°46.24′ E; depth 5509 m; 22 Oct. 2022; H. Nomaki leg.; Dive 1659 of HOV Shinkai 6500, core Red#8; GenBank accession nos: PP662678, PP662679; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr695. The specimen is preserved in a core in 10% buffered formalin.</p><p>Paratype</p><p>NW PACIFIC OCEAN – 32.5° N (west of KEO) • 1 spec.; same data as for holotype; Dive 1659 of HOV Shinkai 6500, core Red#7; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr694. The specimen is preserved in a core in 10% buffered formalin .</p><p>Description</p><p>Overall test morphology</p><p>Holotype (TNS Pr695). In the seabed photograph, the holotype (Fig. 2D) appears as an apparently unstructured cluster of plate-like elements of various shapes and sizes, including one that is fairly large and fan-shaped and others that are more elongate. Most of the main elements are orientated in the same general direction. Shipboard photographs (Supp. file 2: Fig. S4D) show a crooked, elongate, branched structure extending upwards with a small fan-shaped extremity, in addition to plate-like elements, most of which lean outwards away from the centre of the core.</p><p>The preserved specimen includes a main cluster of plate-like elements on one side, and on the other side, a smaller group of plates (not clearly visible in the seafloor image), part of which is attached to a stone (Supp. file 4: µCT video 2; Fig. 9A). A pebble-shaped xenophyophore, described below, partially separates these two groups. The overall appearance is confused and devoid of any obvious pattern. Many of the plates seem rather isolated, possibly because connections are obscured by resedimented material. A μCT render showing the view from above reveals that the main cluster (length 59 mm, width 40 mm), at least, forms a single, complex, inter-connected structure lacking any obvious organisation (Fig. 9A). The other smaller group of plates (length 40 mm, width 15 mm) also seems to have some connection to the main cluster, although this is not entirely clear in the μCT render. The plates have complicated, irregular shapes and are either curved, sometimes strongly, or undulating (Fig. 9B–E; Supp. file 2: Fig. S5E–F; Supp. file 4: µCT video 2). In a few places they are punctuated by one or two small open spaces. The margins are typically lobate, and may give rise to more elongated, rounded, prong-like projections. Other crooked, elongate processes are also developed. The plates are about 1.0– 1.7 mm (mean 1.33 ± 0.20 mm, n = 15) thick.</p><p>Paratype (TNS Pr694). The in situ photograph shows a complex, confusing mass of irregular, poorlyresolved, and apparently plate-like elements, which is difficult to make sense of (Fig. 2C). Shadows suggest that some parts are upstanding. In the shipboard photograph, the most prominent part of the test is a large erect plate with vague concentric zones and an open space at its base (Supp. file 2: Fig. S4C). Adjacent to it is a recumbent plate that is curved into a trough-like structure, behind which two rounded lobes are visible. There is no sign of any symmetry or organisational pattern in either image.</p><p>By the time the paratype was examined in the shore-based lab, much of the test had collapsed. Some parts still projected upwards, but others that were originally upstanding lay flat on the surface of the preserved core (Fig. 10). These collapsed remnants measure 3.9 × 2.5 cm in maximum extent. The main features that remain in what appear to be their original positions are the trough-like plate, which is visible in the µCT renders and shipboard image, and the two rounded lobes (Fig. 10C–F; Supp. file 2: Fig. S5C–D), also seen in the shipboard photograph (Supp. file 2: Fig. S4C–D). These lobes arise from a plate that lies in a vertical plane; they are orientated more of less at right-angles to each other, one projecting vertically, the other horizontally. Some of the other structures visible in the light photographs and µCT scans probably represent parts of the large collapsed plate.</p><p>Surface ornamentation, test structure and composition</p><p>In light photographs, some of the plates display rather irregular concentric zones; these are most apparent on parts of the holotype (Fig. 9D–E) but also visible on the paratype (Fig. 10E; Supp. file 2: Fig. S5C). Radial features, however, have not been observed. Concentric zones with irregular boundaries can be seen in some μCT scans, which show that the internal space within the zones appears to be subdivided further into irregular spaces of different sizes and shapes (Fig. 9B). Micro-CT scans of the surface structure also reveal a rather faint pattern of shallow, cell-like depressions, some of them elongate, across parts of the test surface (Fig. 10B). These features are difficult to interpret. They cannot be seen in light photographs and whether they have any connection with the internal spaces is unclear.</p><p>The test wall is greyish, friable, 210 – 340 µm (mean 295 ± 0.04 µm, n =15) thick and similar to that of Psammina yokosukae sp. nov. When viewed through the wall of the core tube, the test surface in areas not covered in redeposited sediment includes a scattering of larger grains (Supp. file 2: Fig. S5C). Seen under a stereo microscope, the wall is composed of mineral grains of various sizes (Fig. 5A–C). The larger grains are often dark, with a smaller proportion that are whitish and orange.</p><p>Stercomare and granellare</p><p>The stercomare resembles that of Psammina yokosukae sp. nov. in consisting of branches and more irregular masses with varying dimensions that occupy spaces between internal particles and do not conform to any particular trend or pattern. Similarly, the granellare forms narrow strings with a width of 14–40 µm (usually 20–30 µm, mean 24.8 ± 7.26 µm, n = 46). They are closely associated with the internal agglutinated particles and in places are attached to them. The granellare was not observed using SEM, but a strand examined using transmitted light under a high-power microscope did not contain any obvious barite crystals (Fig. 8F–G).</p><p>Molecular characterisation</p><p>Psammina contorta sp. nov. is strongly supported by the BV (98%) and branches at the base of P. tenuis and P. yokosukae sp. nov. The group is supported by 99% BV. The sequenced 18S barcoding fragment of P. contorta sp. nov. contains 1036 nucleotides and the GC content is 36%. The obtained sequences are identical.</p><p>Remarks</p><p>Psammina contorta sp. nov. resembles P. yokosukae sp. nov. in many respects, notably the test structure, the size and composition of the agglutinated grains, and the organisation of the stercomare and granellare systems. However, there are two main morphological differences. First, the test of P. contorta is a highly irregular structure made up of plates of various sizes and shapes that appear somewhat disconnected, in contrast to the more coherent system of fairly well-formed branching plates that characterise the holotype of P. yokosukae . Second, although concentric zones are visible in both light photographs and µCT scans of P. contorta, there is no evidence for internal radial structures running between the zones. Instead, µCT renders appear to show the interior occupied by the irregularly shaped spaces. These rather different morphological characteristics, together with the genetic data, support the recognition of two distinct species.</p><p>As in the case of Psammina yokosukae sp. nov., our phylogenetic reconstruction shows that P. contorta sp. nov. is a close relative of P. tenuis and has similarly narrow granellare branches that are, to some extent, attached to internal test particles. Moreover, the disorganised test is morphologically quite different from the simple curved plate of P. tenuis . As also noted above for P. yokosukae, a particularly striking difference is that the test of P. tenuis is composed largely of radiolarian tests, whereas that of P. contorta consists of mineral grains.</p></div>	https://treatment.plazi.org/id/A5388564FFF5503C5F3322E7FDADFCAA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gooday, Andrew J.;Ishitani, Yoshiyuki;Chen, Chong;Holzmann, Maria;Richirt, Julien;Seike, Koji;Yamashita, Momo;Tsuchiya, Masashi;Nomaki, Hidetaka	Gooday, Andrew J., Ishitani, Yoshiyuki, Chen, Chong, Holzmann, Maria, Richirt, Julien, Seike, Koji, Yamashita, Momo, Tsuchiya, Masashi, Nomaki, Hidetaka (2025): Integrative taxonomy of new xenophyophores (Rhizaria, Foraminifera) from the abyssal northwest Pacific. European Journal of Taxonomy 1004: 144-189, DOI: 10.5852/ejt.2025.1004.2973, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2973/13399
A5388564FFE9503C5F2C20B6FDD2F8AD.text	A5388564FFE9503C5F2C20B6FDD2F8AD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Laminarena Gooday & Ishitani & Chen & Holzmann & Richirt & Seike & Yamashita & Tsuchiya & Nomaki 2025	<div><p>Genus Laminarena gen. nov.</p><p>urn:lsid:zoobank.org:act: 0B9D4CF3-8CC3-49B1-85BB-966EFBBD7C4A</p><p>Type species</p><p>Laminarena variabilis gen. et sp. nov. described below.</p><p>Diagnosis</p><p>Test free, epifaunal, up to at least 6.3 cm maximum dimension. Comprising several thin, delicate, curved or undulating plates with sinuous margins and occasional branches, typically arranged asymmetrically. Obvious apertures absent. Plates marked by concentric zones, traversed by low, closely spaced, radial ridges, together forming distinctive surface ornamentation. Plate walls composed of mineral grains including larger dark particles set in lighter-coloured, fine-grained matrix. Test interior without internal particles except for partitions corresponding to external radial ridges. Dark stercomare branches and pale granellare strands (width 45–100 µm) follow same radial trend. Granellare containing numerous barite crystals.</p><p>Etymology</p><p>A combination of the Latin words ‘ laminam ’ (‘plate’) and ‘ arena ’ (‘sand’), referring to the basically plate-like agglutinated test. Gender feminine.</p><p>Remarks</p><p>Phylogenetically, Laminarena gen. nov. is close to the genus Aschemonella Brady, 1879, grouping together with A. aspera Gooday &amp; Holzmann, 2017 and Aschemonella sp. 3 sensu Gooday et al. (2017b). However, the two genera have little in common morphologically, apart from the fact that the test is agglutinated and internal xenophyae absent. The new genus is characterised by a basically plate-like test, whereas in Aschemonella, the test is either a tube interrupted by internal partitions or a sequence of more or less globular chambers. Given these striking morphological differences, we feel justified in establishing this new genus.</p></div>	https://treatment.plazi.org/id/A5388564FFE9503C5F2C20B6FDD2F8AD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gooday, Andrew J.;Ishitani, Yoshiyuki;Chen, Chong;Holzmann, Maria;Richirt, Julien;Seike, Koji;Yamashita, Momo;Tsuchiya, Masashi;Nomaki, Hidetaka	Gooday, Andrew J., Ishitani, Yoshiyuki, Chen, Chong, Holzmann, Maria, Richirt, Julien, Seike, Koji, Yamashita, Momo, Tsuchiya, Masashi, Nomaki, Hidetaka (2025): Integrative taxonomy of new xenophyophores (Rhizaria, Foraminifera) from the abyssal northwest Pacific. European Journal of Taxonomy 1004: 144-189, DOI: 10.5852/ejt.2025.1004.2973, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2973/13399
A5388564FFEE500D5F762219FBDBF9D2.text	A5388564FFEE500D5F762219FBDBF9D2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Laminarena variabilis Gooday & Ishitani & Chen & Holzmann & Richirt & Seike & Yamashita & Tsuchiya & Nomaki 2025	<div><p>Laminarena variabilis gen. et sp. nov.</p><p>urn: lsid:zoobank.org:act: A7159F47-DD82-4C93-A3A4-D26481268FF2</p><p>Figs 11–19, 21A–E, 22A; Supp. file 2: Figs S6–S 14; Supp. files 5, 6; µCT videos 3–4</p><p>‘ Xenophyophores ’ – Tsuchiya &amp; Nomaki 2021: figs 1–3.</p><p>Diagnosis</p><p>As for genus.</p><p>Etymology</p><p>Latin, ‘ variō ’ (‘diverse or variable’) + suffix ‘- bilis ’, adjective, referring to the variable morphology.</p><p>Material examined</p><p>Holotype NW PACIFIC OCEAN – 30° N • 30°09.2′ N, 143°35.1′ E; depth 5366 m; 23 Oct. 2022; H. Nomaki leg.; Dive 1660 of HOV Shinkai 6500, core Blue#2; GenBank accession no.: PP662670; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr700. The specimen is preserved in a core in 10% buffered formalin.</p><p>Paratype</p><p>NW PACIFIC OCEAN – 30° N • 1 spec.; same data as for holotype; Dive 1660 of HOV Shinkai 6500, core Blue#1; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr699. The specimen is preserved in a core in 10% buffered formalin .</p><p>Other material examined</p><p>NW PACIFIC OCEAN – 30° N • 1 spec.; same data as for holotype; Dive 1660 of HOV Shinkai 6500, core Blue#6; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr702 • 1 spec.; same data as for holotype; Dive 1660 of HOV Shinkai 6500, core Blue#7; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr703 • 1 spec.; same data as for holotype; Dive 1660 of HOV Shinkai 6500, core Red#4; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr705. – 32.5 °N • 1 spec.; 32°34.8′ N, 143°46.2′ E, depth 5505 m; 22 Oct. 2022; H. Nomaki leg.; Dive 1659 of HOV Shinkai 6500, core Red#9; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr697 • 1 spec.; 32°34.7′ N, 143°46.1′ E; depth 5505 m; 24 May 2022; H. Nomaki leg.; Dive 1633 of HOV Shinkai 6500, core Blue#1; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr690 • 1 spec.; 32°34.9′ N, 143°45.9′ E; depth 5505 m; 27 May 2023; H. Nomaki leg.; Dive 1691 of HOV Shinkai 6500, core Blue#0; National Museum of Nature and Science, Tsukuba, Japan, reg. no. TNS Pr714. Specimens are preserved in cores in 10% buffered formalin .</p><p>Description of the typical 30° N form</p><p>Overall test morphology</p><p>The test comprises a more or less complex, asymmetrical system of thin, interconnected plates that follow a curved, or undulating course. Although the plates may appear to some extent separate, in fact they are either parts of one longer, highly sinuous plate or linked to other plates at branching points. There is considerable variation in test morphology and each specimen is different. Micro-µCT scans show that the tests do not penetrate the sediment to any extent and there is no evidence for root-like structures.</p><p>Holotype (TNS Pr700). The seabed photograph of the holotype (Supp. file 2: Fig. S6A) shows two relatively simple, curved plates with the concave sides facing outwards, and another more complex branching plate, all of them displaying concentric zones. In µCT renders, the recovered specimen measures 49 mm by 39 mm in maximum extent and projects to a height of 24 mm above the sediment surface (Fig. 11A, D, F, H). The view from above shows the three main plate-like components that are visible in the seafloor image (Fig. 11A). The plates are thin and delicate, ranging in thickness from 1.2–1.5 mm (mean 1.35 ± 0.12 mm, n = 15). They appear somewhat separated but the μCT video (Supp. file 5) shows that all three arise from a common basal area. Two are relatively simple plates facing at approximately 90 degrees to each other. The largest is strongly curved and forms an almost complete circuit, the second less strongly curved. The third main element branches to create a more complex undulating structure with a sinuous upper margin when viewed from above (Fig. 11A). One part is attached for a short distance to a stone (Fig. 11C–F) and Supp. file 5 appears to show that the largest plate is also attached to a stone. It is possible that the part of the test immediately adjacent the former stone represents the base from which the rest of the test developed.</p><p>Paratype (TNS Pr699). The seabed image (Supp. file 2: Fig. S6B) is dominated by a large, unevenly curved plate with the concave side facing outwards, behind which several additional sinuously curved plates are visible. The recovered specimen is contained within a roughly hemispherical envelope. In µCT renders (Figs 12A, C, E, 13A–C) it measures 63 mm by 58 mm in maximum extent and projects to a height of 42 mm above the sediment surface, making it the largest of the six specimens. The plates range in thickness from 0.60 to 0.85 mm (mean 0.72± 0.09 mm, n = 13). The test appears to lie directly on the sediment surface with no obvious structures penetrating the sediment (Supp. file 6: µCT video 4). The view from directly above (Fig. 13A) reveals a strongly asymmetrical morphology. The plates are located mainly on one side of an open space through which the sediment is visible, with the largest plate dominating the view and partly obscuring the rest of the test. Other µCT renders and light images (Figs 12, 13B–C) show a series of smaller, often strongly curved plates with sinuous rims. Several small, curved plates near the base of the test form tunnel-like features at or just above the sediment surface. There are no reticulations, although the video (Supp. file 6) clearly shows several points at which the plates branch (see also Fig. 12A, C–D). It is also clear from the video that large sections of the test, including the largest plate and others that appear separate from it in some views, in fact comprise a single continuous and highly sinuous plate (Fig. 13B).</p><p>The test has no current connection with several small stones that lie nearby in the core (Fig. 13A). However, careful examination of the µCT renders (Fig. 13A) suggests that a short broken section of the margin near the base of the largest plate was probably originally attached to one of the stones, which still retains a small fragment of the plate on its surface. As in the holotype, it seems likely that this region near the base of the largest plate represents the initial part of the test.</p><p>Additional specimen reg. no. TNS Pr705. In µCT scans the specimen measures 50 mm by 48 mm in maximum extent and projects to a height of 40 mm above the sediment surface. The seafloor image (Supp. file 2: Fig. S6C), together with laboratory photographs and µCT renders (Supp. file 2: Fig. S7), all show what appears to be one continuous main plate that makes several sinuous, sweeping loops in different planes. The largest loop is directed vertically, creating a broad, funnel-like structure that opens in an upward direction, while in the lower part of the test the loops are orientated horizontally (see particularly Supp. file 2: Fig. S7A, E). There are also several additional curved plates near the base of the structure that possibly arise as branches of the main plate, although obvious branching points cannot be seen.</p><p>Additional specimen reg. no. TNS Pr702. The test appears simpler than others in seafloor images. A strongly curved upper plate creates a funnel-like structure through which the sediment surface is visible (Supp. file 2: Fig. S6D). At least one additional plate can be seen beneath the upper plate. The recovered specimen is located to one side of the core and seems to have slumped slightly. It measures 51 mm by 50 mm in maximum extent, projects to a height of 44 mm, and has a distinctly asymmetrical appearance. The µCT top view render shows two strongly concave plates that face in different directions but are linked together to create a single large plate with a strongly sinuous, S-shaped margin (Supp. file 2: Fig. S8A). In side view, another large plate that is strongly curved on a horizontal plane appears to be continuous with the sinuous upper plate (Supp. file 2: Fig. S8B). On one side it branches to give rise to a smaller plate that is curved in the same direction and at one point is attached to a stone (Supp. file 2: Fig. S8B–D). Both these elements merge together into a gently undulating plate near the base of the test.</p><p>Additional specimen reg. no. TNS Pr703. In the seafloor and shipboard images this large specimen is clearly more complex than others and gives the impression of being made up of a larger number of smaller plates (Supp. file 2: Fig. S6E–F). The recovered specimen measures 60 mm by 55 mm in maximum extent and projects to a height of 35 mm. As in the shipboard image, the test appears to comprise numerous curved plates that are orientated in different directions, but with no clear pattern or obvious reticulations (Supp. file 2: Fig. S9). These elements are all interconnected and it is clear from Supp. file 2: Fig. S9A (top view), that many plates that appear distinct from some angles are actually parts of at least two larger plates. One of these is sinuously folded into an S shape, the other follows a straighter course. In places, the plates clearly branch or give rise to side plates. Many parts of the plate system extend upwards but near the base of the test some are more horizontal with their concave sides facing downwards.</p><p>Surface ornamentation, test structure and composition</p><p>The test surfaces of all specimens display a distinctive pattern of concentric zones that follow the shape of the plate margin and are traversed at right angles by low radial ridges separated by shallow furrows (Figs 12, 14A). These features create a distinctive pattern that in places is strongly developed (Supp. file 2: Fig. S7F). In the holotype, the zones are 1.8–5.6 mm (mean 3.23 ± 1.28 mm, n = 7) wide and the radial ridges spaced 0.46–0.70 mm (mean 0.57 ± 0.07 mm, n = 14) apart. In the paratype, the zones also measure between 1.8 and 5.6 mm (usually 2.3–3.9 mm; mean 3.11 ± 0.84 mm, n = 21) wide and the radial ridges are spaced 0.46–0.74 mm (mean 0.57 ± 0.08 mm, n = 16) apart. One of the plates of additional specimen TNS Pr703 features a number of dome-like surface pustules that can be seen in the µCT render (Supp. file 2: Fig. S9B).</p><p>The test plates are 250–320 µm (mean 0.28 ±0.02 µm, n = 16) thick and comprise two more or less parallel walls. Sections that have broken across the concentric zones (i.e., at right angles to the test margin) show that the walls of later (younger) zones overlap those of the preceding zones (Fig. 14C–D), rather than being a simple continuation of the wall. Sections of the test that have broken across the radial ridges (i.e., parallel to the concentric zones and to the test margin) often show that the interior is interrupted by transverse partitions that define compartments (Fig. 14E). Micro-CT scans (Figs 11–13; Supp. file 2: Figs S7–S 9) confirm that the surface pattern of radial ridges and furrows corresponds to these internal partitions. When test fragments are broken to expose the interior, the partitions can be seen as low, parallel structures composed of agglutinated particles arising from the inner surface of the wall (Fig. 14F). Otherwise, the test interior appears largely devoid of internal particles.</p><p>The wall is greyish in overall colour when dried and composed of a mixture of mineral grains of varying sizes (Figs 14A–B, 15). Most of the larger particles are dark but a few are transparent or whitish, although the density of darker grains is higher in some fragments than in others. SEM images show a jumble of particles, mainly of mineral origin and many resembling fragments of volcanic glass (Fig. 15E–F). There is also a tendency for the grain size to increase from the inner to the outer part of a zone (Fig. 15A–C), thereby creating a contrast in grain size across zone boundaries (Fig. 15C–D).</p><p>Granellare and stercomare</p><p>The granellare strands are generally located between the internal partitions (Fig. 16A). They are whitish to pale yellow, branch occasionally, and 45–97 µm wide (mean 67.4 ± 12.8 µm, n = 34). In the fragments examined, the spaces between the partitions are also occupied by masses of loose stercomata or their degraded remnants. In µCT scans of the paratype, the granellare strands appear as bright threads that correspond to the radial lineations (Fig. 13E–F). They are present throughout much of the test but less well developed near the base and more strongly developed towards one side of the upper part (Fig. 13D). The threads branch to some extent. In places near the margin, they appear to be discontinuous across the boundary between the two outer concentric zones (Fig. 13F). In addition to the main radial trend of the granellare, strands running horizontally are visible near the base of some of the concentric zones. Viewed by SEM, fragments of granellare are packed with crystals, a few microns (&lt;5 µm) in size and identical to the barite particles (‘granellae’) that are typical of xenophyophores (Fig. 16C–D). Some crystals are marked by deep, sometimes rectangular depressions.</p><p>Description of 32.5° N form</p><p>This form is represented by two specimens from which we obtained molecular sequences, one collected in 2022 during dive 1659 (TNS Pr697) and the other in 2023 during dive 1691 (TNS Pr714). A third morphologically similar specimen was collected in 2022 during dive 1633 (TNS Pr690) but was immediately preserved in formalin and could not be sequenced.</p><p>Overall test morphology</p><p>Specimen reg. no. TNS Pr697. The test comprises a complex system of irregularly shaped but generally plate-like elements that are often somewhat curved (Fig. 17A–F; Supp. file 2: Fig. S10A–B). In µCT scans, the specimen has an overall extent of ~ 4.3 cm and rises ~ 1.8 cm above the sediment surface. The central part seems to be generally flat lying, but around the periphery, a series of complicated, elongate elements, some of which branch, project upwards to varying extents and at various angles. Four or five of these elements are particularly conspicuous. They project mainly to one side, giving the test an asymmetrical appearance, and tend to widen upwards, sometimes with a vaguely fan-like termination. The plates range in thickness from 0.9 to 1.3 mm (mean 1.08± 0.12 mm, n = 15) and are punctuated by a few open spaces. One section of the periphery forms an undulating margin that rises only slightly from the surface. Concentric zones can be seen across many parts of the test surface (Fig. 17F).</p><p>Specimen reg. no. TNS Pr714. This specimen is similar in overall appearance to TNS Pr697. It forms a delicate structure that includes several fan-shaped plates, one of which is larger than the others (Fig. 17G; Supp. file 2: Figs S10D, S 11). The larger plate is connected to two smaller plates; and one of these merges with a fourth plate that develops from a separate stem. This cluster of plates is linked, in turn, to an irregular formation that includes several bar-like components and irregular excrescences; this part cannot be seen clearly through the core tube because it lies on the edge of the core and is partly obscured by redeposited sediment. The largest plate displays concentric zones and indistinct radial ridges (Fig. 17H), features that are clearly evident in the fragment illustrated in Fig. 18C.</p><p>Specimen reg. no. TNS Pr690. The seabed photograph shows a compact cluster of irregular elements, some of them plate-like (Supp. file 2: Fig. S10E). In the shipboard photograph, the largest element terminates in a distinct fan, punctuated by two elongate open spaces, while bar-like structures project out to one side (Supp. file 2: Fig. S10F). In the µCT render showing the top view, the test appears as a chaotic jumble of relatively small, irregular plate-like elements (Supp. file 2: Fig. S12A). It measures 6.1 cm by 5.9 cm in maximum extent and projects 3.1 cm above the sediment surface. The test structure is clearer in side view renders that show a number of elongate processes that widen to varying extents towards their extremities (Supp. file 2: Fig. S12B–C). These include the fan visible in the shipboard photograph. This and many of the other test parts are orientated upwards at an angle of 40° or more, and are clustered around a prominent, straight, chimney-like tube. However, there are also some narrow structures near the base of the test that extend outwards at a lower angle. In a few places, the plates are perforated by open spaces. The chimney-like tube is probably that of a polychaete. In the shipboard photograph, the upper part of this tube is obscured by a mass of detritus (Supp. file 2: Fig. S10F).</p><p>Surface ornamentation, test structure and composition.</p><p>Although the surface ornamentation that is a feature of the 30° N specimens is less evident in the 32.5° N form, corresponding internal features are obvious in the µCT images of specimen TNS Pr697 (Fig. 17B– C) and particularly specimen TNS Pr690 (Supp. file 2: Fig. S12B–E). These show a distinct pattern of concentric zones and radial structures, presumably internal partitions. The zones are 1.5–4.1 mm (mean 2.88 ± 0.81 mm) wide and the radial features spaced 0.50–1.22 mm (mean 0.87 ± 0.19 mm, n = 17) apart. Partitions are seen in broken cross sections of the specimen TNS Pr714 fragment viewed under a stereo microscope but their development is variable, ranging from vaguely defined to well defined (Fig. 18D–E). Partitions appeared weakly developed on inner test surfaces when this fragment was broken open and viewed under a light microscope (Fig. 18F–G) but were rather more obvious in an SEM image (Fig. 19A).</p><p>In specimens TNS Pr697 and particularly Pr714, the test wall resembles that of the holotype and paratype (Fig. 18A–C; Supp. file 2: Fig. S13A). It is composed of mineral particles of varying sizes, with a matrix of small grains intermingled with a subordinate number of larger grains, mainly whitish and black but occasionally orange. Many of the whitish ones are probably volcanic glass (Supp. file 2: Fig. S13B). The largest are a few hundred microns in size but most are much smaller.</p><p>Granellare and stercomare</p><p>The dissected fragment of specimen TNS Pr714 contained stercomare and granellare, both in good condition (Fig. 18F–G). The stercomare formed several discrete, dark-grey, sausage-shaped masses situated between indistinct ridges that would have formed incomplete radial partitions when the specimen was intact. The masses have the following dimensions: lengths 0.63–2.20 mm (mean 1.23 ± 0.65 mm, n = 6), width 0.20–0.45 mm (mean 0.35 ± 0.07 mm, n = 10), with one longer, branched strand (length at least 3.9 mm), the broken end of which extends beyond the edge of the fragment. The individual stercomata are generally between 8 and 20 µm (mean 14.1 ± 3.24 µm, n = 10) in size (Supp. file 2: Fig. S13C–D). The granellare branches are whitish and lay alongside the stercomare for much of their length (Fig. 18F–G). The main branches are 40–98 µm (mean 65.8 ± 14.9 µm, n = 18) wide; in places, they appear to be closely associated with the stercomare (Fig. 18H). Stercomare masses and granellare branches with similar characteristics were present in a partially dissected fragment of specimen TNS Pr697 (Fig. 18I).</p><p>SEM images of granellare from specimen TNS Pr714 show that intracellular barite crystals are present (Fig. 19C–F). They have a generally oval, sometimes hexagonal morphology and resemble the crystals seen in the 30° N form. Their abundance appears to be quite variable along a granellare strand, with dense concentrations in a few places, sparser numbers or a virtual absence of crystals elsewhere. The dense concentrations are associated with areas where the granellare tube had apparently ruptured. In contrast, only a few could be seen where the tube appeared intact and was strongly crinkled, presumably an artifact of drying (Fig. 19E). However, the crystals probably appear to be sparse in such areas only because our view of the cytoplasm is obscured by the granellare tube wall.</p><p>Granellare fragments from specimens TNS Pr697 were also observed by SEM (Fig. 19G–H). Unfortunately, because they had not been washed adequately, many parts were obscured by salt crystals. Where the granellare surface could be seen clearly, there was no sign of obvious barite crystals within the cytoplasm, although it was not possible to establish whether they were absent or present but obscured by the granellare tube.</p><p>Molecular characterisation</p><p>Laminarena variabilis gen. et sp. nov. is supported by 89% BV and branches as sister to a clade containing representatives of the genera Aschemonella, Abyssalia Gooday &amp; Holzmann, 2020, Psammina, Galatheammina Tendal, 1972 and Moanammina Gooday &amp; Holzmann, 2020 . The branching of the latter clade with L. variabilis is not supported by BV. The sequenced 18S barcoding fragment of L. variabilis contains 1013–1015 nucleotides and the GC content ranges from 36% to 37%.</p><p>Remarks</p><p>The undescribed xenophyophore specimens that were the subject of Tsuchiya &amp; Nomaki’s (2021) feeding experiments came from the same 30° N site as the typical form of Laminarena variabilis gen. et sp. nov. described here and clearly belong to the same morphospecies. A remarkably similar morphotype was photographed (but not collected) in the western CCZ, 7362 km to the southeast of our sampling site (Gooday et al. 2020b: fig. 1f therein). Kamenskaya et al. (2013: fig. 5c, f therein) published in situ photographs of similar specimens from the central CCZ. Probably the morphologically closest described species is Reticulammina plicata Gooday, 1996 from 4613 m depth on the Cape Verde abyssal plain in the NE Atlantic (Gooday 1996). The unique specimen consists of the thin, undulating plates with sinuous margins and several branches although unlike those of the new species, a few wide reticulations are also present. Concentric zones are visible and broken plate edges show signs of partitions that traverse the zones at right angles, although these appear to have no surface expression. Some rather similar but undescribed xenophyophores have been collected on east Pacific seamounts (e.g., Levin et al. 1986: fig. 1e therein; Levin &amp; Thomas 1988: fig. 2c therein).</p><p>In terms of test morphology, the 32.5° N form has a more disorganised appearance compared to the rather elegant typical form from 30° N. Instead of broad, sinuous plates, the test comprises much slenderer elements that radiate outwards, widening to varying extents but mainly towards their upper ends. They sometimes branch and a few are punctuated by open spaces. The surface pattern of concentric zones and radial lineations is much less obvious than in the typical form. However, to some extent, specimen TNS Pr703 from 30° N bridges the morphological gap between the two morphotypes. This specimen has a more complicated test than others from the type locality and appears rather more similar to the 32.5° N form, particularly when the seafloor photographs (Supp. file 2: Figs S6E, S 10A, C, E) are compared.</p><p>Despite their morphological differences, genetic data support the placement of the two forms in the same species. The molecular distance of partial 18S rRNA sequences between specimens from 30° N and 32.5° N is &lt;0.0089 in the Jukes-Cantor model (Jukes &amp; Cantor 1969), and the foraminifera-specific 37f hypervariable regions are almost identical (&gt;99 %), categorizing them as the same species according to the criteria for foraminiferal meta-18S that classifies sequences diverging by less than 1% as conspecifics (Nguyen et al. 2023). Since there has been some diversification, we conducted an Approximately Unbiased test for phylogenetic tree selection (Shimodaira 2002). However, with only three sequenced specimens available to analyse, this failed to reveal any evolutionary scenario (data not shown). More information is clearly required in order to resolve phylogenetic relationships within this species.</p><p>It is not obvious why the 30° N and 32.5° N forms have diverged so much in terms of morphology, or whether there are any environmental drivers that are responsible for the differences between them. As noted above, bottom topography is similar between the two areas, but some environmental differences have been detected (Nomaki unpubl. data). At the 30° N site, CN ratios were higher, and carbon isotopic compositions (δ 13 C) were lower than at 32.5° N, suggesting that the sedimentary organic matter was probably more refractory and may have originated from a different source. This may explain the lower prokaryotic cell numbers and porewater [NH] + concentrations at 30° N compared to 32.5° N, despite TOC concentrations being comparable or even higher. If food is of lower quality at the more southerly site, this would be consistent with the test morphology of the typical form of Laminarena variabilis gen. et sp. nov., which seems better adapted for particle trapping than the more northerly form. Further analysis is needed in order to understand the factors responsible for the striking morphological differences between these two genetically similar xenophyophore populations.</p><p>There are precedents for morphological differences between genetically similar populations of xenophyophores that are separated geographically. Two species from the western CCZ, Aschemonella monilis Gooday &amp; Holzmann, 2017 and Moanammina semicircularis Gooday &amp; Holzmann, 2017, yielded sequences identical to those obtained from specimens collected in the eastern CCZ (Gooday et al. 2020a), although in these cases the geographical separation was 3800 km, compared to only 270 km between the two forms of Laminarena variabilis gen. et sp. nov.</p></div>	https://treatment.plazi.org/id/A5388564FFEE500D5F762219FBDBF9D2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gooday, Andrew J.;Ishitani, Yoshiyuki;Chen, Chong;Holzmann, Maria;Richirt, Julien;Seike, Koji;Yamashita, Momo;Tsuchiya, Masashi;Nomaki, Hidetaka	Gooday, Andrew J., Ishitani, Yoshiyuki, Chen, Chong, Holzmann, Maria, Richirt, Julien, Seike, Koji, Yamashita, Momo, Tsuchiya, Masashi, Nomaki, Hidetaka (2025): Integrative taxonomy of new xenophyophores (Rhizaria, Foraminifera) from the abyssal northwest Pacific. European Journal of Taxonomy 1004: 144-189, DOI: 10.5852/ejt.2025.1004.2973, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2973/13399
