Abstract
A new species and genus of stem-group ‘soft corals’ (Anthozoa: Octocorallia: Alcyonacea) is described and illustrated in detail, as well as compared to other species of the sparse octocorallian fossil record. Sueciatractos leipnitzae gen. et sp. nov. has been collected from the Upper Silurian Hemse beds of the Isle of Gotland, Sweden. The new taxon differs from all other taxa by its unique form of sclerites which were fused at first glance, but stacked compactly, densely packed aggregates forming a nearly solid skeleton or former supporting layers in part. Sueciatractos is compared with other fossil octocoral species that were known by microscopic/mesoscopic sclerites.
Similar content being viewed by others
Avoid common mistakes on your manuscript.
Introduction
Silurian times witnessed one of the most concentrated intervals of reefal development in earth’s history, based in large part on tabulate and rugose coral as well as stromatoporoid assemblages (Zapalski and Berkowski 2019). However, the Palaeozoic/Mesozoic fossil record of other anthozoan cnidarians, such as octocorals, is more than sparse (e.g. Reich 2007, 2009; Fernández-Martínez et al. 2018, 2019). In particular, pre-Ordovician octocorallian taxa are heavily debated (Echmatocrinus—Ausich and Babcock 1998; Reich 2007, 2009 versus Sprinkle and Collins 1998; Pywackia—Taylor et al. 2013 versus Landing et al. 2015), which is in contrast to the time-calibrated trees published recently by Quattrini et al. (2020) and McFadden et al. (2021), where the origin of octocorals was placed in the Ediacaran period. In addition, just recently also the group of Devonian/Carboniferous Heterocorallia has been discussed as putative Octocorallia (Berkowski et al. 2021).
The Silurian octocoral Atractosella (with three named species), although previously misidentified as sponge scleres (Hinde 1887, 1888; Bengtson 1979), echinoid spines (Regnéll 1956) and ostracoderm fish (Lamont 1978), is widespread and has been found in England (Hinde 1887, 1888), Scotland (Stewart et al. 2007; Candela and Crighton 2019), Sweden (Regnéll 1956; Bengtson 1979, 1981a, b; Brood 1981, 1982), Estonia (Hints et al. 2022) and the bottom of the Baltic Sea (Reich 2000, 2002).
Other Palaeozoic octocorallian anthozoans include Nonnegorgonides ziegleri (Dapingian and Sandbian; Lindström 1971) and Petilavenula varifurcata (Floian; Cope 2005) as well as the recently published Catenatus argentinus (Floian–Darriwilian; Carrera et al. 2021), Termieralcyon copperi (Silurian–Permian; Fernández-Martínez et al. 2019), and Lafustalcyon vachardi (Serpukhovian; Denayer et al. 2022).
The aim of this contribution is to report and describe new Silurian octocoral findings, which complement the previously patchy fossil record.
Geological setting, material and methods
Gotland, situated in the Baltic Sea east of the Swedish coast, consists of Silurian rocks ranging from Telychian to Ludfordian strata (latest Llandovery to latest Ludlow) and are dominated by limestones and marls (e.g. Sandström 1998; Sandström et al. 2021; Fig. 1). Reefs and reefal structures on and around Gotland have been mapped and investigated by several researchers (e.g. Hadding 1941; Manten 1971; Eriksson and Laufeld 1978; Bjerkéus and Eriksson 2001; Flodén et al. 2001; Sandström et al. 2021).
The exact regional and stratigraphic reference data for the Tänglings locality (Fig. 1) can be found in the corresponding papers from Laufeld (1974), Larsson (1979), Stridsberg (1985), Bergman (1989), Copper (2004) and Jeppsson et al. (2006). Tänglings is part of the inland central Hemse reef complex which is unfortunately understudied. The Hemse Group is one of the least understood units of the Gotland Silurian sequence (Sandström et al. 2021). After a transgressive phase around the Wenlock–Ludlow boundary with deposition of marls and little or no reef development, a highstand which favoured extensive reef growth with fringing reefs and stromatoporoid biostromes followed (Sandström et al. 2021).
The Tänglings locality contains a rich associated fauna of tabulate and rugose corals, stromatoporoids, echinoids, cyclocystoids, brachiopods, trilobites, ostracods, polychaetes, nautiloids, bryozoans, and sponges (Rothpletz 1913; Regnéll 1956; Martinsson 1962; Ramsköld 1983; Bergman 1995; Stridsberg and Turek 1997; Rhebergen 2005; Young and Kershaw 2005; Kutscher 2008; Reich and Kutscher 2010; Holmer et al. 2013) as well as crinoids, paracrinoids, blastozoans, ophiuroids, asteroids, and gastropods (herein).
All studied octocoral specimens were isolated and picked from residues after washing and sieving unlithified reefal rock samples in hot water or tenside solution, using standard techniques (Wissing et al. 1999). The material is derived from samples collected by Manfred Kutscher (Sassnitz), Heilwig Leipnitz (Uelzen), and Mike Reich. The figured and type material is deposited in the State Natural History Museum, Brunswick (Staatliches Naturhistorisches Museum, Braunschweig), Germany. Additional material will be stored at the Swedish Museum of Natural History (Naturhistoriska riksmuseet), Stockholm, Sweden.
Selected specimens were documented by digital microscopy (Keyence VHX 7000) first, and later coated with gold and studied and photographed using a Phenom XL G2 (ThermoFisher Scientific) scanning electron microscope (SEM) at Brunswick and Munich. The energy-dispersive X-ray spectroscopy (EDS) was also done at the latter SEM.
Institutional abbreviations
SNHMB, Staatliches Naturhistorisches Museum, Braunschweig, Germany; NRM, Naturhistoriska riksmuseet, Stockholm, Sweden.
Systematic palaeontology
The current classification of octocorals is mainly based on the papers and monographs by Wilhelm G. Kükenthal, Sydney J. Hickson and Frederick M. Bayer (see Pérez et al. 2016). Not all of the groups included within the Octocorallia have well-defined synapomorphies, which is why the phylogenies available so far (e.g. Berntson et al. 2001; Won et al. 2001; Sánchez et al. 2003; Park et al. 2012; McFadden et al. 2006, 2021) are not always fully conclusive. However, McFadden et al. (2022) recently revised the higher-level systematics of octocorals, which we follow here.
For terminology of octocoral skeletons and gross morphology used in this study, see Bayer (1956), Bayer et al. (1983), Williams (1993) and Fabricius and Alderslade (2001).
Sub-Phylum Anthozoa Ehrenberg, 1834
Class Octocorallia Haeckel, 1866
Order Malacalcyonacea McFadden, van Ofwegen and Quattrini, 2022
Stem-group ‘Alcyoniina’ Lamouroux, 1812
Genus Sueciatractos nov.
ZooBank LSID. zoobank.org:act:FE717877-0696-4503-A22D-6685D4DE606D.
Type species. Sueciatractos leipnitzae gen. et sp. nov.
Etymology. The generic name is a combination of Latin Suecia (Sweden) and Greek άτρακτος (spindle), referring to the morphology and the origin of the material.
Diagnosis. Colony growth form and shape of polyps unknown. Taxon probably without skeletal axis. Distinct sclerites of coenenchyme are uniform pointed spindles, somewhat angular (Figs. 2C, 4A–G) and regular stacked (Figs. 2C, 5A–D). These densely packed sclerite aggregates may therefore be part of the former supporting layer.
Remarks and discussion. These specimens show a number of distinctive features, which support the erection of a new genus. Sueciatractos gen. nov. differs from all other Palaeozoic alcyonacean (sclerite-bearing) genera (Atractosella, Figs. 2A, B, 3A–K; Catenatus; Lafustalcyon) by the stacked arrangement of sclerites (Fig. 5A–D) as well as by the form and shape of these sclerites (Fig. 4A–G). Only the sclerites of Termieralcyon are comparably densely packed as aggregates. However, the Termieralcyon sclerite morphotypes described by Fernández-Martínez et al. (2019: fig. 5) are quite distinct in morphology and size from the Sueciatractos sclerites described here.
Range. Silurian: Ludlow; Gotland, Sweden.
Sueciatractos leipnitzae sp. nov.
ZooBank LSID. zoobank.org:act:A25FD6FC-7BB8-4AFA-8B10-0AD86AEFB821.
Material. 102 single sclerites and 71 stacks of sclerites (SNHMB-G.5511–5672), including the holotype (SNHMB-G.5500, Fig. 4A) and paratypes (SNHMB-G.5501, 5502, 5505–5509, Figs. 4B, C, F, G, 5A–C) as well as further figured material (SNHMB-G.5503, 5504, 5510, Figs. 4D, E, 5D).
Locality and horizon. Tänglings, Gotland, Sweden (see Laufeld 1974: 133; Larsson 1979: 179; Stridsberg 1985: 63; Bergman 1989: 127; Copper 2004: 156 and Jeppsson et al. 2006: fig. 1). From the lower part of the Late Silurian Hemse Group; Ludlow: latest Gorstian (or the middle part = ‘Etelhem Fm.’; Ludlow: earliest Ludfordian).
Etymology. The species epithet honours Heilwig Leipnitz (*1928; Uelzen, Germany) for her generous palaeontological donations to public museums (and in honour of her 95th birthday this year).
Diagnosis. As for the genus.
Description. Morphology of soft parts like colony shape, branching patterns, polyps, etc., unknown. Coenenchyme sclerites are simple, straight, uniform, spindle-shaped on both sides, and somewhat angular in longitudinal direction (length 1.0–4.0 mm). The sclerites are thickest in the middle, with a diameter of about 0.45–1.0 mm. No obvious, or only slight/smooth surface ornamentation of sclerites (due to the diagenetic overprint; cf. also Majoran 1987), visible (Fig. 4A–G). Sclerites are densely packed aggregates (Figs. 2C, 5A–D) probably of the former inner or outer supporting layer.
All collected sclerites and stacks of sclerites are preserved in the form of low-magnesium calcite (chemical composition identified by EDS analysis). In contrast, modern alcyoniid sclerites are composed of high-magnesium calcite (cf. Conci et al. 2021). However, the preservation of our fossil material as low-magnesium calcite is typical and not surprising due to diagenetic overprinting.
Remarks and discussion. The Telychian/Sheinwoodian/Homerian Atractosella cataractaca Bengtson, which is stratigraphically and regionally close, has only loosely embedded sclerites (Fig. 2A, B) that show a typical surface granulation (Figs. 2B, 3A–K) and appear not to be irregularly cemented (like in Pleistocene/Holocene “Sinularia spiculites” or “Alcyonarian limestones”; cf. Konishi 1981; Accordi et al. 1989; Schuhmacher 1997; Jeng et al. 2011; Fig. 2F). The morphological variability of the sclerites (bent and branched/multiaxial spindles; Figs. 2B, 3F, K) is also clearly greater in Atractosella than in Sueciatractos gen. nov. Modern revisions are lacking for the other two Atractosella species—the Telychian A. andreae (Lamont) and the Homerian A. siluriensis Hinde. However, these seem very similar to A. cataractaca described from Gotland, Sweden (Bengtson 1981b), the Baltic Sea bedrock (Reich 2002) and Estonia (Hints et al. 2022). The Serpukhovian Lafustalcyon vachardi Denayer et al. and the Silurian–Permian Termieralcyon copperi Fernández-Martínez et al. clearly show a different morphology and ornamentation of sclerites, which are not fused, but densely packed (Fernández-Martínez et al. 2019; Denayer et al. 2022).
Modern octocoral species with densely packed sclerite aggregates are known from various clades of the Scleralcyonacea and Malacalcyonacea (see McFadden et al. 2022), whereas species with fused sclerites are known from members of the Clavulariidae, Alcyoniidae and Xeniidae (all Malacalcyonacea), among others (e.g. Bayer 1981, 1995; van Ofwegen and Haddad 2011; Halász et al. 2014).
Discussion
Interpretation of octocoral fossils is often challenging, due to differences in the range of characters (sclerites, axes, etc.) available for study. Nearly all octocorals possess microscopic sclerites, which are found in varying concentrations embedded in the coenenchyme. These calcitic bodies, providing support and protection, around 5.0–0.02 mm in size, are the most important feature used in the identification of octocorals (e.g. Bayer 1956; Bayer et al. 1983; Fabricius and Alderslade 2001). Also, the microstructure as well as the arrangement of the sclerites in the octocoral colonies can be an important taxonomic feature (Aharonovich and Benayahu 2012; Fabricius and Alderslade 2001; McFadden et al. 2022). In addition, there is a co-occurrence of different sclerite types within one species and colony (e.g. Tentori and van Ofwegen 2011). Therefore, it is often difficult to assign isolated fossil sclerites to specific higher-level groups within the Alcyonacea, and other Octocorallia. Our new genus and species described here (Sueciatractos leipnitzae) can undoubtedly be assigned to the ‘Alcyoniina’ (or stem-group ‘Alcyoniina’), since the absence of an axis and simple spindle-shaped sclerites are synapomorphies of these.
However, the octocorallian fossil record known so far is very patchy (Reich 2007, 2009). In addition to the Palaeozoic taxa already discussed, there is also Mesozoic and Cenozoic evidence. Isolated alcyonacean sclerites (formerly of the Alcyoniina, Holaxonia, Stolonifera, Scleraxonia, Calcaxonia; now Malacalcyonacea and Scleralcyonacea) were mostly recorded from Cretaceous (Počta 1886; Papp 1972; Alexandrowicz 1977; Moosleitner 1990; Herrig et al. 1996; Reich and Frenzel 2002; Reich et al. 2005; Schlagintweit and Gawlick 2009; Frenzel et al. 2014), Paleogene (Hickson 1938; Deflandre-Rigaud 1955, 1956, 1957; Kocurko 1988; Kocurko and Kocurko 1992) and Neogene (Deflandre-Rigaud 1955, 1956, 1957; Kristan-Tollmann 1966; Langer 1989) sediments. Jurassic reports (Hasse 1890; Cayeux 1921) are, unfortunately, still questionable, even if partially articulated octocorals are known from similar strata (Barthel 1978; Heyng and Viohl 2015). Only a few partially articulated finds or larger parts (e.g. holdfasts, axes) of octocorals are also known from later geological periods (e.g. Voigt 1958; Giammona and Stanton jr. 1980; Kocurko 1988; Kocurko and Kocurko 1992; Bayer 1992; Helm and Schülke 2003; Lozouet and Molodtsova 2008; Zuschin and Gebhardt 2009; Löser 2000, 2016).
However, the Permo–Triassic gap of this record is certainly striking. To what extent this is related to the different seawater chemistry (calcite versus aragonite seas) in earth’s history remains to be clarified in the future.
Conclusions
This study increases our knowledge on the early evolutionary history and diversification of soft corals and their relatives. Our results show that the Alcyoniina (or stem-group Alcyoniina) was already present with various soft coral skeletal structures in Silurian times, regardless of the large gaps of articulated/isolated alcyonacean sclerite material in the later fossil record.
This group of octocorals complements and increases the biodiversity of stromatoporoid–tabulate/rugose coral associations (and the Anthozoa in general) within the Silurian ecosystems of Gotland, Sweden.
In addition, we continue to believe that the incomplete alcyonacean fossil record is primarily related to examination gaps and the limited number of octocoral palaeontologists, rather than preservational gaps. However, as the primary biomineralization of octocorals is also an important factor of their diversity, more basic and advanced studies on the octocorallian fossil record are urgently needed.
Availability of data and material
All data are provided in the text and the figures.
References
Accordi, G., F. Carbone, and R. Matteucci. 1989. “Alcyonarian spiculite” nei calcari recifali Quaternari della costa Somala. Rendiconti della Società Geologica Italiana 12: 17–20.
Aharonovich, D., and Y. Benayahu. 2012. Microstructure of octocoral sclerites for diagnosis of taxonomic features. Marine Biodiversity 42: 173–177. https://doi.org/10.1007/s12526-011-0102-3.
Alexandrowicz, S.W. 1977. Sclerites of octocorals from the Upper Cretaceous of eastern Poland. Journal of Paleontology 51 (4): 687–692.
Ausich, W.I., and L.E. Babcock. 1998. The phylogenetic position of Echmatocrinus brachiatus, a probable octocoral from the Burgess Shale. Palaeontology 41 (2): 193–202.
Barthel, K.W. 1978. Solnhofen. Ein Blick in die Erdgeschichte, 1–393. Thun: Ott.
Bayer, F.M. 1956. Octocorallia. In Treatise on Invertebrate Paleontology, Part F, Coelenterata, ed. R.C. Moore, F166–F231. New York, N.Y.: Geological Society of America and Lawrence, Kans.: University of Kansas Press.
Bayer, F.M. 1981. On some genera of stoloniferous octocorals (Coelenterata: Anthozoa), with descriptions of new taxa. Proceedings of the Biological Society of Washington 94 (3): 878–901.
Bayer, F.M. 1992. The Helioporacean Octocoral Epiphaxum, Recent and Fossil. A Monographic Iconography. Studies in Tropical Oceanography, Miami 15: i–vii + 1–76.
Bayer, F.M. 1995. Two new species of the alcyonacean genus Protodendron (Octocorallia: Alcyoniidae) from the Indian Ocean off Natal. Bulletin of Marine Science 57 (2): 301–312.
Bayer, F.M., M. Grasshoff, and J. Verseveldt, eds. 1983. Illustrated trilingual glossary of morphological and anatomical terms applied to Octocorallia, 1–75. Leiden: E.J. Brill / Dr. W. Backhuys.
Bengtson, S. 1979. Sponges. In Lower Wenlock Faunal and Floral Dynamics—Vattenfallet Section, Gotland, eds. V. Jaanusson, S. Laufeld, and R. Skoglund. Sveriges Geologiska Undersökning (C: Avhandlingar och uppsatser) 762: 61–62. (=SGU Årsbok 73(3)).
Bengtson, S. 1981a. En läderkorall i Gotlands silur. Fauna och Flora 76 (1): 37–42.
Bengtson, S. 1981b. Atractosella, a Silurian alcyonacean octocoral. Journal of Paleontology 55 (2): 281–294.
Bergman, C.F. 1989. Silurian paulinitid polychaetes from Gotland. Fossils and Strata 25: 1–128.
Bergman, C.F. 1995. Symmetroprion spatiosus (Hinde), a jawed polychaete showing preference for reef environments in the Silurian of Gotland. GFF 117 (3): 143–150. https://doi.org/10.1080/11035899509546210.
Berkowski, B., M.K. Zapalski, E. Jarochowska, and P. Alderslade. 2021. Early development and coloniality in Oligophylloides from the Devonian of Morocco—are Heterocorallia Palaeozoic octocorals? PLoS ONE 16 (9): e0257523 [1-25]. https://doi.org/10.1371/journal.pone.0257523.
Berntson, E.A., F.M. Bayer, A.G. McArthur, and S.C. France. 2001. Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rRNA sequences. Marine Biology 138: 235–246. https://doi.org/10.1007/s002270000457.
Bjerkéus, M., and M. Eriksson. 2001. Late Silurian reef development in the Baltic Sea. GFF 123 (3): 169–179. https://doi.org/10.1080/11035890101233169.
Brood, K. 1981. Svepelektronmikroskopet—ett genombrott för mikropaleontologin. Fauna och Flora 76 (6): 253–258.
Brood, K. 1982. Gotländska fossil, 1–95. Stockholm: P.A. Norstedt & Söners Förlag.
Candela, Y., and W.R.B. Crighton. 2019. Synoptic revision of the Silurian fauna from the Pentland Hills, Scotland described by Lamont (1978). Palaeontologia Electronica 22.2.19A: 1–45. https://doi.org/10.26879/868.
Carrera, M.G., G.G. Voldman, M.J. Mango, and G.P. Nestell. 2021. Ordovician enigmatic sclerite-type elements from western Argentina: possible oldest axial components of alcyonacean octocorals. Acta Palaeontologica Polonica 66 (3): 535–544. https://doi.org/10.4202/app.00869.2020.
Cayeux, L. 1921. Existence de nombreux spicules d’Alcyonaires dans les minerais de fer jurassique de France. Comptes rendus hebdomadaires des séances de l‘Académie des Sciences 172 (16): 987–988.
Conci, N., S. Vargas, and G. Wörheide. 2021. The biology and evolution of Calcite and Aragonite Mineralization in Octocorallia. Frontiers in Ecology and Evolution 9: 623774 [1–15]. https://doi.org/10.3389/fevo.2021.623774.
Cope, J.C.W. 2005. Octocorallian and hydroid fossils from the Lower Ordovician of Wales. Palaeontology 48 (2): 433–445. https://doi.org/10.1111/j.1475-4983.2005.00455.x.
Copper, P. 2004. Silurian (Late Llandovery–Ludlow) Atrypid Brachiopods from Gotland, Sweden, and the Welsh Borderlands, Great Britain, i–x + 1–215. Ottawa: NRC Research Press. https://doi.org/10.1139/9780660190112.
Deflandre-Rigaud, M. 1955. Sur les sclérites d’Alcyonaires fossils et leur classification Micralcyonarites manip. Nov. du Miocène moyen d’Australie. Comptes rendus des séances de l‘Académie des Sciences 241 (19): 1327–1329.
Deflandre-Rigaud, M. 1956. Les sclérites d’Alcyonaires fossiles. Elements d’une classification. Annales de Paléontologie 42: 3–24.
Deflandre-Rigaud, M. 1957. A classification of fossil alcyonarian sclerites. Micropaleontology 3 (4): 357–366. https://doi.org/10.2307/1484441.
Denayer, J., E. Poty, F. Tourneur, and M. Aretz. 2022. Colonial Heterocorallia (Cnidaria, Anthozoa) and their epibionts from the lower Carboniferous of Montagne Noire and Pyrenees, southern France. PalZ. Paläontologische Zeitschrift. https://doi.org/10.1007/s12542-021-00588-1.
Ehrenberg, C.G. 1834. Beiträge zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin (Physikalische Klasse) 1832 (1): 225–380.
Eriksson, C.-O., and S. Laufeld. 1978. Philip structures in the submarine Silurian of northwest Gotland. Sveriges Geologiska Undersökning (C: Avhandlingar och uppsatser) 736: 1–30 (=SGU Årsbok 71(11)).
Fabricius, K.K., and P.P. Alderslade. 2001. Soft Corals and Sea Fans: A comprehensive guide to the tropical shallow-water genera of the Central-West Pacific, the Indian Ocean and the Red Sea, 1–264. Townsville: AIMS.
Fernández-Martínez, E., I. Coronado, S. Rodríguez, and F. Tourneur. 2018. Hallazgo de escleritos de Alcionáceos en materiales Paleozoicos: desentramando el registro de Octocorallia. In Yacimientos paleontológicos excepcionales en la península Ibérica, eds. N. Vaz, and A.A. Sá. Cuadernos del Museo Geominero 27: 571–578.
Fernández-Martínez, E., I. Coronado, S. Rodríguez, F. Tourneur, and M. Badpa. 2019. Alcyonacea awakens: Palaeobiology and palaeoecology of Palaeozoic octocorals known from their sclerites. Geological Journal 54 (6): 3593–3618. https://doi.org/10.1002/gj.3359.
Flodén, T., M. Bjerkéus, I. Tuuling, and M. Eriksson. 2001. A Silurian reefal succession in the Gotland area. GFF 123 (3): 137–152. https://doi.org/10.1080/11035890101233137.
Frenzel, P., M. Reich, and E. Herrig. 2014. Ein Meer am Ende des Mesozoikums. Kreide. In Lebensspuren im Stein: Ausflüge in die Erdgeschichte Mitteleuropas, eds. P. Rothe, V. Storch, and C. von See, 166–176. Weinheim: Wiley-VCH Verlag.
Giammona, C.P., and R.J. Stanton. 1980. Octocorals from the Middle Eocene Stone City Formation, Texas. Journal of Paleontology 54 (1): 71–80.
Hadding, A. 1941. The Pre-Quaternary sedimentary rocks of Sweden. VI. Reef limestones. Lunds Universitets Årsskrift (N.F., Avdelningen 2) 37 (10): 1–137 (=Kungliga Fysiografiska Sällskapet i Lund handlingar (N.F.) 52(10)).
Haeckel, E. 1866. Generelle Morphologie der Organismen. Zweiter Band: Allgemeine Entwickelungsgeschichte der Organismen, i–clx + 1–462. Berlin: G. Reimer.
Halász, A., C.S. McFadden, D. Aharonovich, R. Toonen, and Y. Benayahu. 2014. A revision of the octocoral genus Ovabunda Alderslade, 2001 (Anthozoa, Octocorallia, Xeniidae). ZooKeys 373: 1–41. https://doi.org/10.3897/zookeys.373.651.
Hasse, C. 1890. Fossile Alcyonarien. Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie 1890 (II): 59–65.
Helm, C., and I. Schülke. 2003. An almost complete specimen of the Late Cretaceous (Campanian) octocoral ‘Isis’ ramosa Voigt (Gorgonacea) from the Lower Saxony Basin, northwest Germany. Cretaceous Research 24 (1): 35–40. https://doi.org/10.1016/S0195-6671(03)00020-X.
Herrig, E., H. Nestler, P. Frenzel, and M. Reich. 1996. Discontinuity Surfaces in the high Upper Cretaceous of Northeastern Germany and their Reflection by Fossil Associations. In Global and Regional Controls on Biogenic Sedimentation. II. Cretaceous Sedimentation. Research Reports. Göttinger Arbeiten zur Geologie und Paläontologie Sb3: 107–111.
Heyng, A.M., and G. Viohl. 2015. Octokorallen (Octocorallia). In Solnhofen. Ein Fenster in die Jurazeit. Volume 1, eds. G. Arratia, H.-P. Schultze, H. Tischlinger, and G. Viohl, 204–205. München: F. Pfeil.
Hickson, S.J. 1938. An alcyonarian from the Eocene of Mississippi. Journal of the Washington Academy of Sciences 28 (2): 49–51.
Hinde, J.J. 1887. A Monograph of the British Fossil Sponges. Part 1. Palaeontographical Society, Monographs 40 (for 1886): 1–92.
Hinde, J.J. 1888. A Monograph of the British Fossil Sponges. Part 2. Palaeontographical Society, Monographs 41 (for 1887): 93–188.
Hints, L., H. Pärnaste, P. Männik, M. Reich, and S. Rozhnov. 2022. Development of faunal diversity during the late Llandovery—early Wenlock in the easternmost part of the Baltic Palaeobasin—implications for the Ireviken Event. Estonian Journal of Earth Sciences 71 (2): 89–110. https://doi.org/10.3176/earth.2022.07.
Holmer, L.E., L. Popov, and M.G. Bassett. 2013. Silurian craniide brachiopods from Gotland. Palaeontology 56 (5): 1029–1044. https://doi.org/10.1111/pala.12033.
Jeng, M.-S., H.-D. Huang, C.-F. Dai, Y.-C. Hsiao, and Y. Benayahu. 2011. Sclerite calcification and reef-building in the fleshy octocoral genus Sinularia (Octocorallia: Alcyonacea). Coral Reefs 30: 925–933. https://doi.org/10.1007/s00338-011-0765-z.
Jeppsson, L., M.E. Eriksson, and M. Calner. 2006. A latest Llandovery to latest Ludlow high-resolution biostratigraphy based on the Silurian of Gotland—a summary. GFF 128 (2): 109–114. https://doi.org/10.1080/11035890601282109.
Kocurko, M.J. 1988. Notes on fossil octocorals and comparison of some modern and ancient octocoral remains. Tulane Studies in Geology and Paleontology 21 (3–4): 105–115.
Kocurko, M.J., and D.J. Kocurko. 1992. Fossil Octocorallia of the Red Bluff Formation, lower Oligocene, Mississippi. Journal of Paleontology 66 (4): 594–602. https://doi.org/10.1017/S0022336000024458.
Konishi, K. 1981. Alcyonarian spiculite: limestone of soft corals. In Proceedings of the Fourth International Coral Reef Symposium, Volume 1, 643 – 649. Manila: Marine Science Center, University of the Philippines.
Kristan-Tollmann, E. 1966. Alcyonarien-Sklerite aus dem Torton des Burgenlandes, Österreich. Sitzungsberichte der Österreichischen Akademie der Wissenschaften (Mathematisch-naturwissenschaftliche Klasse, Abteilung 1) 175 (4–6): 129–141.
Kutscher, M. 2008. Papillicalymene sinuata n. sp., eine neue Trilobiten-Art aus dem Silur von Gotland (Schweden). Geschiebekunde aktuell 24 (4): 129–134.
Lamont, A. 1978. Pentlandian miscellany: Mollusca, Trilobita, etc. Scottish Journal of Sciences 1 (5): 245–302.
Lamouroux, J.V.F. 1812. Extrait d’un mémoire sur la classification des Polypiers coralligènes non entierement pierreux. Nouveau Bulletin des Sciences (par la Société philomatique de Paris) 3 (63): 181–188.
Landing, E., J.B. Antcliffe, M.D. Brasier, and A.B. English. 2015. Distinguishing Earth’s oldest known bryozoan (Pywackia, late Cambrian) from pennatulacean octocorals (Mesozoic–Recent). Journal of Paleontology 89 (2): 292–317. https://doi.org/10.1017/jpa.2014.26.
Langer, M. 1989. Haftorgan, Internodien und Sklerite von Keratoisis melitensis (Goldfuss, 1826) (Octocorallia) in den pliozänen Foraminiferenmergeln („Trubi“) von Milazzo (Sizilien). Paläontologische Zeitschrift 63 (1/2): 15–24. https://doi.org/10.1007/BF02989523.
Larsson, K. 1979. Silurian tentaculitids from Gotland and Scania. Fossils and Strata 11: 1–180.
Laufeld, S. 1974. Reference localities for palaeontology and geology in the Silurian of Gotland. Sveriges Geologiska Undersökning (C: Avhandlingar och uppsatser) 705: 1–172 (=SGU Årsbok 68(12)).
Lindström, M. 1971. An octocoral from the Lower Ordovician of Sweden. Geologica et Palaeontologica 12: 41–52.
Löser, H. 2000. Repertoire of species. Catalogue of Cretaceous Corals 1: 1–135.
Löser, H. 2016. Systematic part. Catalogue of Cretaceous Corals 4: 1–710.
Lozouet, P., and T. Molodtsova. 2008. Filling a gap: the first occurrences of Epiphaxum (Cnidaria: Helioporacea: Lithotelestidae) in the Eocene, Oligocene and Miocene. Palaeontology 51 (1): 241–250. https://doi.org/10.1111/j.1475-4983.2007.00744.x.
Majoran, S. 1987. Structural investigations of octcoral sclerites. Zoologica Scripta 16 (4): 277–287. https://doi.org/10.1111/j.1463-6409.1987.tb00074.x.
Manten, A.A. 1971. Silurian reefs of Gotland. Developments in Sedimentology 13: 1–539.
Martinsson, A. 1962. Ostracodes of the Family Beyrichiidae from the Silurian of Gotland. Bulletin of the Geological Institutions of the University of Uppsala 41: 1–369.
McFadden, C.S., S.C. France, J.A. Sánchez, and P. Alderslade. 2006. A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences. Molecular Phylogenetics and Evolution 41 (3): 513–527. https://doi.org/10.1016/j.ympev.2006.06.010.
McFadden, C.S., A.M. Quattrini, M.R. Brugler, P.F. Cowman, L.F. Dueñas, M.V. Kitahara, D.A. Paz-García, J.D. Reimer, and E. Rodríguez. 2021. Phylogenomics, origin, and diversification of Anthozoans (Phylum Cnidaria). Systematic Biology 70 (4): 635–647. https://doi.org/10.1093/sysbio/syaa103.
McFadden, C.S., L.P. van Ofwegen, and A.M. Quattrini. 2022. Revisionary systematics of Octocorallia (Cnidaria: Anthozoa) guided by phylogenomics. Bulletin of the SSB (society of Systematic Biologists) 1 (3): 8735. https://doi.org/10.18061/bssb.v1i3.8735.
Moosleitner, G. 1990. Lederkorallen aus den alpinen Gosauschichten. Fossilien 7 (5): 206–207.
Ofwegen, L.P., and M.A. Haddad. 2011. A probably invasive new genus and new species of soft coral (Octocorallia: Alcyonacea: Clavulariidae) from Brazil. Zootaxa 3107 (1): 38–46. https://doi.org/10.11646/ZOOTAXA.3107.1.2.
Papp, A. 1972. Foraminiferen und Mikrofazies der Nordzone. In Studien in der Unterkreide des Wienerwaldes, W. Grün, G. Kittler, G. Lauer, A. Papp, W. Schnabel, and O. Čorna. Jahrbuch der geologischen Bundesanstalt Wien 115 (2): 119–122.
Park, E., D.-S. Hwang, J.-S. Lee, J.-I. Song, T.-K. Seo, and Y.-J. Won. 2012. Estimation of divergence times in cnidarian evolution based on mitochondrial protein-coding genes and the fossil record. Molecular Phylogenetics and Evolution 62 (1): 329–345. https://doi.org/10.1016/j.ympev.2011.10.008.
Pérez, C.D., B. de Moura Neves, R.T. Cordeiro, G.C. Williams, and S.D. Cairns. 2016. Diversity and Distribution of Octocorallia. In The Cnidaria, Past, Present and Future, eds. S. Goffredo, and Z. Dubinsky, 109–123. Springer International Publishing Switzerland. https://doi.org/10.1007/978-3-319-31305-4_8
Počta, P. 1886. Über fossile Kalkelemente der Alcyoniden und Holothuriden und verwandte recente Formen. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften (Mathematisch-naturwissenschaftliche Classe) 92 (for 1885) (I): 7–12.
Quattrini, A.M., E. Rodríguez, B.C. Faircloth, P.F. Cowman, M.R. Brugler, G.A. Farfan, M.E. Hellberg, M.V. Kitahara, C.L. Morrison, D.A. Paz-García, J.D. Reimer, and C.S. McFadden. 2020. Palaeoclimate ocean conditions shaped the evolution of corals and their skeletons through deep time. Nature Ecology & Evolution 4: 1531–1538. https://doi.org/10.1038/s41559-020-01291-1.
Ramsköld, L. 1983. Silurian cheirurid trilobites from Gotland. Palaeontology 26 (1): 175–210.
Regnéll, G. 1956. Silurian echinoids from Gotland. Arkiv för Mineralogi och Geologi 2 (7): 155–178.
Reich, M. 2000. Skleren von Oktokorallen aus einem Silur-Geschiebe Vorpommerns. Geschiebekunde aktuell 16 (2): 59–61.
Reich, M. 2002. Skleren von Alcyonacea (Anthozoa: Octocorallia) aus einem Silur-Geschiebe Norddeutschlands. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2002 (9): 551–561. https://doi.org/10.1127/njgpm/2002/2002/551.
Reich, M. 2007. Where are all the fossil octocorals (Cnidaria: Anthozoa)? In The Palaeontological Association, 51st Annual Meeting, 16th-19th December 2007, Uppsala University, Sweden. Abstracts, ed. Anonymous. The Palaeontological Association Newsletter 66: 89.
Reich, M. 2009. A critical review of the octocorallian fossil record (Cnidaria: Anthozoa). In International Conference on the Cambrian Explosion – Walcott 2009, eds. M.R. Smith, L.J. O’Brien, and J.-B. Caron, 85. Toronto: ROM.
Reich, M., and P. Frenzel. 2002. Die Fauna und Flora der Rügener Schreibkreide (Maastrichtium, Ostsee). Archiv für Geschiebekunde 3 (2/4): 73–284.
Reich, M., and M. Kutscher. 2010. Cyclocystoids (Echinodermata: Echinozoa) from the Silurian of Gotland, Sweden. In Echinoderms: Durham, eds. L.G. Harris, S.A. Böttger, C.W. Walker, and M.P. Lesser, 67–70. London etc.: Taylor & Francis. https://doi.org/10.1201/9780203869543-c11
Reich, M., P. Frenzel, and E. Herrig. 2005. Ein Meer am Ende der Oberkreide. Die Schreibkreide. Biologie in unserer Zeit 35 (4): 260–267. https://doi.org/10.1002/biuz.200410285.
Rhebergen, F. 2005. Sponges (Porifera) from Silurian strata on Gotland. GFF 127 (3): 211–216. https://doi.org/10.1080/11035890501273211.
Rothpletz, A. 1913. Über die Kalkalgen, Spongiostromen und einige andere Fossilien aus dem Obersilur Gottlands. Sveriges Geologiska Undersökning (Ca: Afhandlingar och uppsatser) 10: 1–57.
Sánchez, J.A., H.R. Lasker, and D.J. Taylor. 2003. Phylogenetic analyses among octocorals (Cnidaria): mitochondrial and nuclear DNA sequences (lsu-rRNA, 16S and ssu-rRNA, 18S) support two convergent clades of branching gorgonians. Molecular Phylogenetics and Evolution 29 (1): 31–42. https://doi.org/10.1016/S1055-7903(03)00090-3.
Sandström, O. 1998. Sediments and stromatoporoid morphotypes in Ludfordian (Upper Silurian) reefal sea stacks on Gotland, Sweden. GFF 120 (4): 365–371. https://doi.org/10.1080/11035899801204365.
Sandström, O., P. Dahlqvist, M. Erlström, L. Persson, S. Kershaw, and M. Calner. 2021. Stratigraphy of the Gorstian and Ludfordian (upper Silurian) hemse Group reefs on Gotland, Sweden. GFF 143 (1): 71–83. https://doi.org/10.1080/11035897.2020.1858959.
Schlagintweit, F., and H.-J. Gawlick. 2009. The incertae sedis Carpathoporella Dragastan, 1995, from the Lower Cretaceous of Albania: skeletal elements (sclerites, internodes/branches, holdfasts) of colonial octocorals. Facies 55: 553–573. https://doi.org/10.1007/s10347-009-0185-5.
Schuhmacher, H. 1997. Soft corals as reef builders. In Proceedings of the 8th International Coral Reef Symposium, Panama, June 24–29, 1996. Vol. 2, eds. H.A. Lessios, and I.G. Macintyre, 499–502. Balboa: Smithsonian Tropical Research Institute.
Sprinkle, J., and D. Collins. 1998. Revision of Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia. Lethaia 31 (4): 269–282. https://doi.org/10.1111/j.1502-3931.1998.tb00517.x.
Stewart, S.E., L.I. Anderson, and E.N.K. Clarkson. 2007. Miscellanea. In Silurian fossils of the Pentland Hills, Scotland, eds. E.N.K. Clarkson, D.A.T. Harper, C.M. Taylor, and L.I. Anderson, 195. London: The Palaeontological Association. (=Palaeontological Association, Field Guides to Fossils 11)
Stridsberg, S. 1985. Silurian oncocerid cephalopods from Gotland. Fossils and Strata 18: 1–65.
Stridsberg, S., and V. Turek. 1997. A revision of the Silurian nautiloid genus Ophioceras Barrande. GFF 119 (1): 21–36. https://doi.org/10.1080/11035899709546450.
Taylor, P.D., B. Berning, and M.A. Wilson. 2013. Reinterpretation of the Cambrian ‘bryozoan’ Pywackia as an octocoral. Journal of Paleontology 87 (6): 984–990. https://doi.org/10.1666/13-029.
Tentori, E., and L.P. van Ofwegen. 2011. Patterns of distribution of calcite crystals in soft corals sclerites. Journal of Morphology 272 (5): 614–628. https://doi.org/10.1002/jmor.10942.
Voigt, E. 1958. Untersuchungen an Oktokorallen aus der oberen Kreide. Mitteilungen aus dem Geologischen Staatsinstitut in Hamburg 27: 5–49.
Williams, G.C. 1993. Coral Reef Octocorals: An Illustrated Guide to the Soft Corals, Sea Fans and Sea Pens inhabiting the Coral Reefs of Northern Natal, 1–63. Durban: Natural Science Museum.
Wissing, F.-N., E. Herrig, and M. Reich. 1999. Arbeitstechniken der Mikropaläontologie. Eine Einführung, 1–191. Stuttgart: F. Enke.
Won, J.H., B.J. Rho, and J.I. Son. 2001. A phylogenetic study of the Anthozoa (phylum Cnidaria) based on morphological and molecular characters. Coral Reefs 20: 39–50. https://doi.org/10.1007/s003380000132.
Young, G.A., and S. Kershaw. 2005. Classification and controls of internal banding in Palaeozoic stromatoporoids and colonial corals. Palaeontology 48 (3): 623–651. https://doi.org/10.1111/j.1475-4983.2005.00480.x.
Zapalski, M.K., and B. Berkowski. 2019. The Silurian mesophotic coral ecosystem: 430 million years of photosymbiosis. Coral Reefs 38 (1): 137–147. https://doi.org/10.1007/s00338-018-01761-w.
Zuschin, M., and H. Gebhardt. 2009. Octocorals as hosts to serpulid-macroids from the Cretaceous of the Potiguar Basin, Brazil. Lethaia 42 (3): 381–382. https://doi.org/10.1111/j.1502-3931.2009.00176.x
Acknowledgements
Our paper is dedicated to Hans-Georg Herbig on occasion of his 65th birthday and official retirement from the chair of palaeontology at Cologne University. We thank two anonymous reviewers and the editor Hannes Löser (Hermosillo, México) for their constructive comments as well as Michael Amler (Cologne, Germany) for the invitation to contribute to this special issue. In addition, we would like to extend our gratitude to Heilwig Leipnitz, Uelzen, Germany, for her donations of specimens, discussion and other support during the last three decades.
Funding
Open Access funding enabled and organized by Projekt DEAL.
Author information
Authors and Affiliations
Corresponding author
Additional information
Handling Editor: Hannes Loeser.
This work is registered in ZooBank, the LSID is: urn:lsid:zoobank.org:pub:D129C6AA-7C09-4CC4-8AEF-C5581894A669.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Reich, M., Kutscher, M. A new alcyonacean octocoral (Anthozoa) from the Late Silurian of Gotland, Sweden. PalZ 97, 729–739 (2023). https://doi.org/10.1007/s12542-023-00654-w
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12542-023-00654-w