Early Badenian transgression on the outer flank of Western Carpathian Foredeep, Hluchov area, Czech Republic

This multidisciplinary study, based on borehole cores from the Hluchov area in Czech Republic, docu- ments an early Badenian marine transgression on the outer flank of the Western Carpathian Foredeep . The shallow-marine deposits represent coastal transgression over a terrestrial topography of weathered pre-Cenozoic bedrock. The lower facies association (FA1) consists of siliciclastic sediment derived from local substrate erosion. Facies indicate a wave-dominated environment with unstable bottom, variable rate of sediment supply and an incremental rise of relative sea level. The upper facies association (FA2) consists of carbonates indicating a major landward shift of shoreline, decline in siliciclastic input and further sea-level rise. The succession represents a transgressive to highstand systems tract. The maximum flooding surface, ca . 1 m above the FA1/FA2 boundary, is signified by an anomalous decrease in K and Th, an increased Th/K ratio and highest U concentration. The heavy-mineral assemblages in FA1 confirm local sediment provenance, whereas those in FA2 indicate broader sediment derivation, including volcanic component from contemporaneous rhyolitic to rhyodacitic eruptions. The deposits contain a wide range of marine fauna, with the foraminifers and molluscs indicating an early Badenian age. Molluscs, bryozoans and echinoderms indicate a normal-salinity environment with a decreasing hydraulic energy. Foraminifers indicate salinity fluctua- tions in the lowest part of the succession. The isotopic composition of mollusc shells shows marked inter-species differences and a general negative shift in the d 13 C and d 18 O values, indicating diagenetic alteration. The impact of diagenetic processes appears to have been controlled by sedimentary facies. The highly negative d 13 C and d 18 O values correspond to sediment layers with the highest Th/K ratios and hence low clay content. Sediment permea- bility was thus probably crucial in controlling the differential circulation and impact of diagenetic fluids

Paleozoic origins of cheilostome bryozoans and their parental care inferred by a new genome-skimmed phylogeny

Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic

Cyclostome bryozoans from Qingdao, South Yellow Sea, China

Liu, H., Liu, X., Zágoršek, K. (2019): Cyclostome bryozoans from Qingdao, South Yellow Sea, China. Zootaxa 4603 (3): 473-500, DOI: 10.11646/zootaxa.4603.3.

Hemipustulopora tuoyuan Liu & Liu & Zágoršek 2019, n. sp.

<i>Hemipustulopora tuoyuan</i> n. sp. <p>(Figs 46–49)</p> <p>lsid:zoobank.org:act: D0A1C723-B08E-4F19-B35A-D489DE51CF01</p> <p> <b>Diagnosis.</b> Colony large, autozooids arranged in indistinct transversal rows. Gonozooid very large, extending over up to 20 autozooidal tubes, frontal wall indistinct. Ooeciopore elongated to circular, attached to the autozooidal tube. Pseudopores large, numerous, often close by radially arranged laminae.</p> <p> <b>Holotype.</b> Specimen MBM 284503 from Jiaozhou Bay (Figs 46–48).</p> <p> <b>Paratypes.</b> Specimens MBM 284360, MBM 284358 (Jiaozhou Bay) and MBM 284445 (Lingshan Island).</p> <p> <b>Additional material.</b> More than 10 colonies from samples MBM 194701, MBM 0 92324, MBM 284363, MBM 284361, MBM 284385, MBM 194797, MBM 0 92236 and MBM 194840 (Jiaozhou Bay), MBM 194797, MBM 0 92370 (Lingshan Island), MBM 284502 from Nanhai road.</p> <p> <b>Etymology.</b> Referring to the ellipsoidal shape of the ooeciopore. In Chinese phonetic transcription, ‘ellipsoidal’ is <i>tuǒyuán</i>.</p> <p> <b>Description.</b> Colony erect, lobate, large, branches up to 2 mm wide, distance between bifurcations up to 3.5 mm. Autozooids densely arranged, in somewhat transverse rows, peristomes short, apertures only on one side, almost circular, approximately 160–180 µm in diameter. Pseudopores large (diameter approximately 13–15 µm) closed by radially arranged laminae (Fig. 48). Gonozooid very large, perforated by autozooidal tubes, frontal wall only slightly elevated; pseudopores the same as that of autozooidal tubes (diameter approximately 9–10 µm), and therefore, the gonozooid almost indistinct. Ooeciopore elongate, oval to elliptical (up to 210 µm wide by 80–100 µm long), rarely circular (diameter of approximately 120 µm). Ooeciostome slightly turned towards the direction of growth, attached to one autozooidal tube. Protoecium not observed. Dorsal side of the colony with distinct, concentric growth lines and minute pseudopores.</p> <p> <b>Remarks.</b> The species is similar to <i>Entalophora multipora</i> MacGillivray, 1895 (p. 140, pl. 21, fig. 1) in the general shape and size of the colony and the arrangement of autozooid. However, no gonozooids have been described in the latter species. <i>Diastopora patina</i> Lamarck as described by MacGillivray (1895, p. 130, pl. 20, figs 4, 5) has a similar arrangement of autozooidal tubes, but differs in having much smaller gonozooids, which are not perforated by autozooidal tubes.</p> <p> The species belongs to <i>Hemipustulopora</i> because of the large colonies with wide branches and apertures only on one side, arranged in slightly transverse rows; frontal, much extended gonozooid; and smooth dorsal side with visible, concentric growth lines. <i>Hemipustulopora</i> has been recorded only from South Africa (Brood 1976).</p> <p> <i>Filisparsa</i> d’Orbigny, 1853 is another very similar genus in having apertures only on one side and a smooth back side of the branches. The type species of <i>Filisparsa</i> is, however, Early Cretaceous and has rather narrow, nonflabellate branches.</p> <p>A few specimens (MBM 284502 from Nanhai Road) have circular ooeciopore (diameter ca. 120 µm), with ooeciostome directed upwards. This type of ooeciopore is situated close to the neighbouring autozooidal tube, but usually not directly attached to it. We consider this difference as within species variability.</p> <p> <b>Occurrence.</b> Nanhai Road, Jiaozhou Bay and Lingshan Island.</p>Published as part of <i>Liu, H., Liu, X. & Zágoršek, K., 2019, Cyclostome bryozoans from Qingdao, South Yellow Sea, China, pp. 473-500 in Zootaxa 4603 (3)</i> on pages 491-493, DOI: 10.11646/zootaxa.4603.3.3, <a href="http://zenodo.org/record/2702283">http://zenodo.org/record/2702283</a&gt

Idmidronea erecta Liu & Liu & Zágoršek 2019, n. sp.

Idmidronea erecta n. sp. (Figs 22–25) lsid:zoobank.org:act: 190A0C00-38AB-43B1-B1D8-6060329257E3 Diagnosis. Idmidronea with gonozooid frontally situated between the fascicles, and straight ooeciostome arranged between regular autozooidal apertures. Holotype. Specimen MBM 284448 from Lingshan Island (Figs 22, 23). Paratype. Specimen MBM 284471 (Jiaozhou Bay) Additional material. Two colonies without gonozooecium from samples MBM 0 92370 (Lingshan Island) and MBM 0 92378 (Jiaozhou Bay). Etymology. Referring to the straight direction of the ooeciostome, not curved as commonly in other Idmidronea species. Description. Colony erect, robust, and dichotomously branched. Branches short and wide (length about 1000–1300 µm, width at the proximal end of the branch about 670–820 µm), with ca. 5–8 zooidal fascicles between successive bifurcations. Fascicles alternating, consist of 3–6 autozooidal apertures. Autozooidal tubes wide (ca. 138–153 µm); length of the frontal wall 380–456 µm, diameter of apertures ca. 100 µm. Gonozooid frontal, elongate (length about 1300 µm), spreading over three or four fascicles, frontal wall porous. Ooeciopore oval, slightly smaller than the autozooidal aperture (diameter approximately 70–80 µm), situated close to the bifurcation, proximally oriented, usually close to a regular autozooid. Dorsal side of the branch porous, almost flat and slightly convex. Kenozooids approximately 40 µm in diameter, developed on the proximal portion of the branch, spreading over the colony in the distal direction. Remarks. Idmidronea atlantica as described by Hayward & Ryland (1985) is similar to the new species in the position of the gonozooid, but differs in having much longer branches and the swan-neck-shaped ooeciostome. Idmidronea biporata Brood, 1976 also has a straight ooeciostome but differs in having very delicate colonies, much narrower branches, and fascicles consisting of only two autozooidal apertures. Many recent Idmidronea species have a similar arrangement of autozooidal tubes, and similar-sized branches and pseudopores, but lack a gonozooid to compare with the new species. For example, Idmonea contorta Busk, 1875 (p. 12, pl. 8) from South Africa shows similar arrangement of autozooidal tubes as does Idmonea australis MacGillivray, 1885b (p. 470, pl. 68, fig. 2). In addition to the presence of kenozooids, which may be a variable character that may (or may not) develop in basal branches during late astogeny, this species differs from Exidmonea intercalata n. sp. in having straight oriented ooeciostome, situated more or less in the central part of the gonozooid. Exidmonea intercalata n. sp. has downwardly curved, swan-neck shaped ooeciostome, situated in the margin of the gonozooid. Occurrence. Lingshan Island and Jiaozhou Bay.Published as part of Liu, H., Liu, X. & Zágoršek, K., 2019, Cyclostome bryozoans from Qingdao, South Yellow Sea, China, pp. 473-500 in Zootaxa 4603 (3) on pages 482-483, DOI: 10.11646/zootaxa.4603.3.3, http://zenodo.org/record/270228

Tubulipora perforata Liu & Liu & Zágoršek 2019, n. sp.

<i>Tubulipora perforata</i> n. sp. <p>(Figs 11–15)</p> <p>lsid:zoobank.org:act: 612DC68B-D2A8-4263-AFF7-9CCF73C62C38</p> <p> v. <i>Tubulipora pulchra</i>: Liu, Yin & Ma, 2001: p. 383, pl. 5, figs 1–3.</p> <p> v. <i>Tubulipora pulchra</i>: Seo, 2005: p. 275, pls 7, 8.</p> <p> <b>Diagnosis.</b> <i>Tubulipora</i> with autozooid arranged in fascicles containing 2–10 autozooidal tubes, protoecium with smooth margin. Gonozooid slightly triangular, with oval ooeciopore adjacent to the proximal or near the proximal margin of the closest distal autozooidal tube. Basal lamina visible beyond the budding zone.</p> <p> <b>Holotype.</b> Specimen MBM 284359 from Jiaozhou Bay (Figs 11, 12).</p> <p> <b>Paratypes.</b> Specimens MBM 284369, MBM 284362, MBM 284490, MBM 284346 (Jiaozhou Bay).</p> <p> <b>Additional material.</b> More than 10 colonies from samples MBM 194708, MBM 194683, MBM 194656, MBM 194700, MBM 194710, MBM 194676 and MBM 194667 (Jiaozhou Bay).</p> <p> <b>Etymology.</b> Referring to the large pseudopores of the gonozooecium.</p> <p> <b>Description.</b> Colony encrusting, almost circular, sometimes lobate. Autozooids wide (approximately 100 µm), arranged in somewhat radial rows. Apertures rounded to square-shaped, often in fascicles of 2–10 during late astogeny. Peristomes long (ca. 160–220 µm), slightly curved and perforated by very small pseudopores (ca. 4 µm in diameter). Budding margin with 2–4 rows of partially developed zooids overlapping the margin, forming a characteristic edge, without the basal lamina limited to the margin of the colony, not extending further. Protoecium approximately 100–110 µm in diameter, often with smooth margins. Gonozooid slightly triangular, longer (1250–1660 µm) than wide (620–1100 µm), spreading over 2–5 autozooidal tubes and perforated by these tubes; pseudopores numerous, large (diameter approximately 12 µm). Ooeciopore elongated oval, centrally situated, approximately 100–120 µm wide by 40–50 µm long, adjacent to the proximal margin of the succeeding autozooid. Ooeciostome with distinctly extended lips developed around the ooeciopore.</p> <p> <b>Remarks.</b> The new species differs from other <i>Tubulipora</i> species in having autozooidal tubes in fascicles, combined with a very large, elongated, almost triangular gonozooid with elongated oval ooeciopore attached to the proximal margin of the succeeding autozooidal tube, non-serrate protoecium and basal lamina projecting beyond the budding zone.</p> <p> The new species differs from <i>T. similis</i> in having a subcircular, non-serrate protoecium and a somewhat upright ooeciostome (vertical to the colony surface) with slightly flared lips, while in <i>T. similis</i> the ooeciostome is slightly curved. Moreover, the new species has autozooecial apertures that are polygonal, arranged in fascicles and the ooeciopore is elongate oval, while in the <i>T. similis</i> the ooeciopore is circular.</p> <p> <i>Tubulipora concinna</i> is very similar to the new species. MacGillivray’s (1885a, p. 94, pl. 1, fig. 10) illustration shows the projecting basal lamina, autozooidal tubes in fascicles, extended gonozooid and oval ooeciopore. However, it differs in the more lateral position (relative to the following autozooid) of the ooeciopore, and in the gonozooid being wider than long. The original material of MacGillivray (1885a) is not accessible; therefore, the exact attribution of his species remains uncertain. Moreover, <i>T. concinna</i> was synonymised with <i>Diastopora latomarginata</i> d’Orbigny, 1853 by Waters (1889) and consequently also by Jelly (1889).</p> <p> Specimens of <i>T. pulchra</i> in Liu <i>et al.</i> (2001) and Seo (2005) show the same characters as the new species (i.e. large, almost triangular gonozooid with oval ooeciopore attached to the succeeding autozooidal tube). Therefore, we synonymised this material with our new species.</p> <p> <b>Occurrence.</b> Jiaozhou Bay in Qingdao and also Jeju Island in Korea (Seo 2005).</p>Published as part of <i>Liu, H., Liu, X. & Zágoršek, K., 2019, Cyclostome bryozoans from Qingdao, South Yellow Sea, China, pp. 473-500 in Zootaxa 4603 (3)</i> on pages 478-480, DOI: 10.11646/zootaxa.4603.3.3, <a href="http://zenodo.org/record/2702283">http://zenodo.org/record/2702283</a&gt

Exidmonea intercalata Liu & Liu & Zágoršek 2019, n. sp.

Exidmonea intercalata n. sp. (Figs 16–21) lsid:zoobank.org:act: E7C19B17-7B70-45A6-989B-DCC1004FD20D Diagnosis. Exidmonea with 2–6 autozooidal tubes in each fascicle, gonozooid situated at bifurcation, with ooeciopore located on its distalmost edge, downwardly curved, swan-neck shaped, hidden/immersed between bifurcating branches. Holotype. Specimen MBM 284444 from Lingshan Island (Figs 16–18). Paratypes. Specimens MBM 284469, MBM 284470, MBM 284472, MBM 284376 (Jiaozhou Bay) and MBM 284499 (Jiaozhou Bay). Additional material. More than 10 colonies from samples MBM 0 92370, MBM 194661 (Lingshan Island), MBM 0 92378 (Jiaozhou Bay) and MBM 0 91767 (Jiaozhou Bay) Etymology. Referring to the immersed/submerged ooeciostome in the zone of bifurcation. Description. Colony erect, dichotomously branched. Branches short (the distance between successive bifurcations approximately 880–1380 µm) and narrow (430–500 µm wide before bifurcation). Approximately 6–8 autozooidal fascicles between successive branch bifurcations. Fascicles alternating, each consisting of 2–6 autozooidal apertures. Distance between fascicles in the longitudinal direction ca. 210–300 µm. Apertures oval to rounded rectangular, diameter approximately 90–110 µm. Gonozooid triangular, small (ca. 410–460 µm wide by 500–520 µm long), situated frontally in the zone of bifurcation, spreading over two or three fascicles, with densely porous frontal wall (pseudopore diameter ca. 8–10 µm). Ooeciopore located on the distalmost edge of the gonozooid, hidden between bifurcating branches, narrower than autozooidal tubes (ca. 60 µm wide). Ooeciostome short, downwardly curved, swan-neck shaped, adjacent to a deformed autozooidal tube with elongated oval aperture. Dorsal side of the colony flat, with arcuate growth lines and pseudopores arranged parallel to them; kenozooids absent. Remarks. Idmonea pauper Canu & Bassler (1929, p. 545, pl. 84, figs 13, 14) is similar to the new species in the size and position of gonozooid, but differs in having ooeciopore adjacent to the first peristome of a fascicle. The downwardly curved, swan-neck shaped ooeciostome of the new species is similar to that of Idmidronea atlantica (Forbes in Johnston, 1847), as described by Hayward & Ryland (1985). However, I. atlantica differs mainly in having much denser fascicles and longer branches (more than 20 fascicles between successive bifurcation and the branch is more than 4 mm long). Moreover, I. atlantica develops a kenozooidal overgrowth on the dorsal side of the branches. Occurrence. Jiaozhou Bay and Lingshan Island.Published as part of Liu, H., Liu, X. & Zágoršek, K., 2019, Cyclostome bryozoans from Qingdao, South Yellow Sea, China, pp. 473-500 in Zootaxa 4603 (3) on pages 480-481, DOI: 10.11646/zootaxa.4603.3.3, http://zenodo.org/record/270228

FIGURES 43–45 in Cyclostome bryozoans from Qingdao, South Yellow Sea, China

FIGURES 43–45. Nevianipora pulcherrimoidea (Liu in Liu, Yin & Ma, 2001) n. comb. Lingshan Island. 43. MBM 284478, general view showing the arrangement of autozooids. 44, 45. MBM 284335 and MBM 284447, details of two gonozooids with an oval ooeciopore and ooeciostome with extended lips. Scale bars: 43 = 1 mm; 44, 45 = 100 µm