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Brachiopod Phylogeny in the Cambrian Guliforms, Obolellates and Rhynchonelliforms (E.G., Zhang Et Al., 2009, 2014, 2015; Holmer Et Al., 2018A)

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Permophiles Issue #66 Supplement 1

pods which may further address these questions.

Glenn A. Brock

The world’s oceans are changing. IPCC (2013) predictions suggest that by the end of the current century our seas will be

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a challenge to all organisms, how will it impact brachiopods? Given that they have a higher proportion of mineralised tissue than virtually any other invertebrate group what will be the

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instance or repairing and maintaining that shell once it is made? In this talk I will review a series of experiments and historical studies undertaken with Emma Cross and Lloyd Peck (Cross et al., 2015, 2016, 2018) that seek to explore the answers to these questions.
Department of Biological Sciences, Macquarie University, NSW 2109, Australia

Leonid E. Popov

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CF10 3NP, UK
Brachiopods are richly represented in the rock record and as early as the Cambrian, where they show an impressive diversity of form and in shell morphology (e.g., Harper et al., 2017).

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zoan branch of the bilaterian tree based on molecular data. Our

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Cambrian organophosphatic tommotiids as belonging to the brachiopod stem (e.g., Holmer et al., 2002). Subsequent discov-

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Paterimitra, and the inferred bivalved scleritome of Micrina from the lower Cambrian of South Australia reveals these three tom-

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in the Paterimitra includes a preserved pedicle tube (Skovsted et al., 2008, 2009; Holmer et al., 2008, 2011). It is likely that the Brachiopoda may have evolved, by sclerite reduction and modi-

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Micrina and Paterimitra also preserve traces of their earliest ontogeny, including bivalved larval shells with evidence of the earliest larval attachment. The conjoined bivalved shell of adult living brachiopods most likely represents a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan probably evolved independently in living linguliform and rhynchonelliforms (Holmer et al., 2011; see also Zhang et al., 2018a,b). Studies of new records of exceptionally preserved soft-bodied fossil assemblages (Konservat-Lagerstätten) are also critical. For example, we have now been able to reassess highly problematic Chengjiang fossils such as the Cotyledion

References

&URVVꢁꢀ(ꢄꢀ/ꢄꢁꢀ3HFNꢁꢀ/ꢄꢀ6ꢄꢀ ꢀ+DUSHUꢁꢀ(ꢄꢀ0ꢄꢀꢅꢆꢇꢊꢄꢀ2FHDQꢀDFLGL¿FD-

tion does not impact shell growth or repair of the Antarctic brachiopod Liothyrella uva (Broderip, 1833). Journal of

Experimental Marine Biology and Ecology, 462, 29–35.

Cross, E.L., Peck, L.S., Lamare, M.D. & Harper, E.M. 2016.

1RꢀRFHDQꢀDFLGL¿FDWLRQꢀH൵HFWVꢀRQꢀVKHOOꢀJURZWKꢀDQGꢀUHSDLUꢀLQꢀ

the New Zealand brachiopod Calloria inconspicua (Sowerby,

1846). ICES Journal of Marine Sciences, 73, 920-926.

Cross, E. L., Harper, E. M. & Peck, L. S. 2018. A 120-year record of resilience to environmental change in brachiopods. Global Change Biology. ISSN 1354-1013 DOI 10.1111/ gcb.14085
Harper, E.M. & Peck, L.S. 2016. Latitudinal and depth gradients in marine predation pressure. Global Ecology and

Biogeography 25, 670-678.

IPCC. 2013. Climate change 2013: The physical science basis. In
T. F. Stocker, D. Qin, G-K. Plattner, M. Tignor, S. K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex & P. M. Midgley (Eds.),

:RUNLQJꢀJURXSꢀ,ꢀFRQWULEXWLRQꢀWRꢀWKHꢀ¿IWKꢀDVVHVVPHQWꢀUHSRUWꢀRIꢀ

the intergovernmental panel on climate change (pp. 1–1522). Cambridge, United Kingdom and New York, NY, USA: Cambridge University Press.

  • as
  • a
  • sclerite-covered sessile tube-shaped probable stem

“lophophorate” (Zhang et al., 2013). We have also recognized Yuganotheca from the Chengjiang as a tubular, agglutinated stem lophophorate (Zhang et al., 2013). This observation adds to our understanding of more derived brachiopods, such as linguliforms, obolellates and rhynchonelliforms (e.g., Zhang et al., 2009, 2014, 2015; Holmer et al., 2018a). Exceptionally preserved pedunculated Cambrian rhynchonelliforms from both the Chengjiang and Utah also indicate that early members of this most important group of living brachiopods had two widely

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tion of the Brachiopoda in the Cambrian remain problematic and controversial issues. Presently only two of the three major brachiopod clades, the Linguliformea and the Rhynchonelliformea, have known Cambrian representatives. The origin and phylogeny of the Craniiformea remains problematic, but it is possible that Cambrian rhynchonelliform chileate-like brachiopods are involved in the processes.

Plenary lecture

Brachiopod Phylogeny in the Cambrian

Lars E. Holmer

Shaanxi Key laboratory of Early Life and Environments and Department of Geology, State Key Laboratory of Continental Dynamics, Northwest University, Xi’an 710069, China

Department of Earth Sciences, Palaeobiology, Uppsala University, SE-752 36 Uppsala, Sweden

Zhifei Zhang

Shaanxi Key laboratory of Early Life and Environments and Department of Geology, State Key Laboratory of Continental Dynamics, Northwest University, Xi’an 710069, China

Zhiliang Zhang

References

Shaanxi Key laboratory of Early Life and Environments and Department of Geology, State Key Laboratory of Continental Dynamics, Northwest University, Xi’an 710069, China
Harper, D., Popov, L. & Holmer, L.E. 2017. Brachiopods, origin and early history. Palaeontology, 60, 609-631

56

Permophiles Issue #66 Supplement 1

Holmer, L.E., Skovsted, C.B. & Williams, A. 2002. A stem

'LYHUVL¿FDWLRQꢀSDWWHUQVꢀRIꢀEUDFKLRSRGVꢀ

after the end Ordovician mass extinction

group brachiopod from the Lower Cambrian – support for

a Micrina (halkieriid) ancestry. Palaeontology, 45, 875-882.

Holmer, L. E., Skovsted, C. B., Brock, G. A., Valentine, J.L. & DQGꢀLWVꢀSDODHRELRJHRJUDSKLFꢀVLJQL¿FDQFHꢀ
Paterson, J. R. 2008. The Early Cambrian tommotiid Micrina,

Bing Huang

a sessile bivalved stem group brachiopod. Biological letters,
State Key Laboratory of Paleobiology and Stratigraphy, Nanjing
4, 724-728.

Institute of Geology and Palaeontology, The Chinese Academy

of Sciences, Nanjing 210008, PR China
Holmer, L.E., Skovsted, C.B., Larsson, C., Brock, G.A. &
Zhang, Z. 2011. First record of a bivalve larval shell in Early

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Palaeontology, 54, 235-239.

Jisuo Jin

Department of Earth Sciences, University of Western Ontario,
Holmer, L.E., Zhang, Z., Topper, T., Popov, L. & Claybourn, T.
London, Ontario, N6A 5B7, Canada
2018a. The attachment strategies of Cambrian kutorginate

brachiopods, the curious case of two pedicle openings and

WKHLUꢀSK\ORJHQHWLFꢀVLJQL¿FDQFHꢄꢀJournal of Paleontology (in

press).

Jiayu Rong

State Key Laboratory of Paleobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, The Chinese Academy of Sciences, Nanjing 210008, PR China
Holmer, L.E., Popov, L., Ghobadi Pour, M., Claybourn, T.,
Zhang, Z., Brock, G.A. & Zhang, Z. 2018b. Evolutionary sig-

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of the pedunculate rhynchonelliform brachiopod Nisusia sul-

cata. Lethaia (in press).

Skovsted, C, Brock, G, Paterson, J, Holmer, L. & Budd, G 2008
The scleritome of Eccentrotheca from the Lower Cambrian

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for tommotiid phylogeny. Geology, 36, 171-174.
Skovsted, C.B., Holmer, L.E., Larsson, C.M, Högström, A.E.S.,
Brock, G.A., Topper, T.P., Balthasar, U., Petterson Stolk, S. & Paterson, J.R.. 2009. The scleritome of Paterimitra: an early Cambrian stem group brachiopod from South Australia.

Proceedings of the Royal Society B, 276, 1651-1656.

Zhang, Z.F., Li, G.X., Emig, C.C., Han, J., Holmer, L.E. & Shu,
D.G. 2009. Architecture and function of the lophophore in the problematic brachiopod Heliomedusa orienta (Early Cambrian, South China). Geobios, 42: 649-661.
Zhang, Z., Holmer, L.E., Skovsted, C., Brock, G. & Budd, G.
2013. A sclerite-bearing stem group entoproct from the early Cambrian and its implications. 6FLHQWL¿Fꢀ5HSRUWVꢁ 3, 1066.
Zhang, Z.-F., Li, G.-X., Holmer, L.E., Brock, G.A., Balthasar, U.,
Skovsted, C.B., Fu, D-J., Zhang, X.-L., Wang, H.-Z., Butler, A., Zhang, Z.-L., Cao, C-Q., Han, J., Liu, J.-N. & Shu, D.-G. 2014. An early Cambrian agglutinated tubular lophophorate with brachiopod characters. 6FLHQWL¿Fꢀ5HSRUWV, 4682.
Zhang Z.F., Zhang, Z.L., Holmer , L.E. & Li, G.X. 2015. First report of linguloid brachiopods with soft parts from the lower Cambrian (Series 2, Stage 4) of the Three Gorges area, South

China. Annales de Paléontologie, 101:167–177.

The Late Ordovician biotic crisis was associated with a brief but intense glaciation episode in earth history. Post-glacial marine transgression created vast habits in epicontinental seas, in which

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one of the most abundant and diverse fossil groups. Based on detailed data of brachiopod occurrences after the end-Ordovician mass extinction, together with newly published data, we analyzed

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to the Aeronian (early Silurian) by Network Analysis (Fig. 1) and Frequency Distribution Analysis. Rhuddanian brachiopod faunas were characterized by low diversity and localized high abundance, except for a relatively high-diversity early Rhuddanian fauna in Avalonia-Baltica and South China. Invariably, these faunas were predominated by re-established holdover and once-cosmopolitan taxa from the Late Ordovician, primarily orthides and strophomenides. By the Aeronian, global brachiopod diversity nearly doubled in comparison with the Rhuddanian, owing to a major

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tropics and high-latitude Gondwana, as well as the concomitant proliferation of endemic and cosmopolitan taxa, in association with a global expansion of epicontinental seas and heterogeneity of specialized local habitats. A drastic turnover of brachiopods

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Paleozoic evolutionary fauna. With the ordinal data of brachiopods in major palaeoplates, we discussed the turnover process. Compared with Ordovician orthides and strophomenides, which globally increased their diversity after the extinction, atrypides and pentamerides of Silurian group displayed a drastic diversi-

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continents, demonstrating the two new orders preferred more warm water environments than the Ordovician groups. Atrypides

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pentamerides delayed until to Aeronian, and this is typically demonstrated by the record of many endemic taxa from South China. The Silurian-type atrypide and pentameride communities thrived mainly in relatively shallow environments (BA2–3) during the Rhuddanian, but expanded and dominated in deeper, mid-shelf and outer-shelf settings (BA3–5) by the Aeronian (Huang et al., 2018).
Zhang, Z., Zhang, Z., Holmer, L.E. & Chen, F. 2018a. Postmetamorphic allometry in the earliest acrotretoid brachiopods from the lower Cambrian (Series 2) of South China, and its implications. Palaeontology, 61, 183–207.
Zhang, Z., Popov, L., Holmer, L.E. & Zhang, Z. 2018b. Earliest

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evidence for metamorphosis and its implications. BMC evo-

lutionary biology (in press).

57

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  • International Chronostratigraphic Chart

    International Chronostratigraphic Chart

    INTERNATIONAL CHRONOSTRATIGRAPHIC CHART www.stratigraphy.org International Commission on Stratigraphy v 2014/02 numerical numerical numerical Eonothem numerical Series / Epoch Stage / Age Series / Epoch Stage / Age Series / Epoch Stage / Age Erathem / Era System / Period GSSP GSSP age (Ma) GSSP GSSA EonothemErathem / Eon System / Era / Period EonothemErathem / Eon System/ Era / Period age (Ma) EonothemErathem / Eon System/ Era / Period age (Ma) / Eon GSSP age (Ma) present ~ 145.0 358.9 ± 0.4 ~ 541.0 ±1.0 Holocene Ediacaran 0.0117 Tithonian Upper 152.1 ±0.9 Famennian ~ 635 0.126 Upper Kimmeridgian Neo- Cryogenian Middle 157.3 ±1.0 Upper proterozoic Pleistocene 0.781 372.2 ±1.6 850 Calabrian Oxfordian Tonian 1.80 163.5 ±1.0 Frasnian 1000 Callovian 166.1 ±1.2 Quaternary Gelasian 2.58 382.7 ±1.6 Stenian Bathonian 168.3 ±1.3 Piacenzian Middle Bajocian Givetian 1200 Pliocene 3.600 170.3 ±1.4 Middle 387.7 ±0.8 Meso- Zanclean Aalenian proterozoic Ectasian 5.333 174.1 ±1.0 Eifelian 1400 Messinian Jurassic 393.3 ±1.2 7.246 Toarcian Calymmian Tortonian 182.7 ±0.7 Emsian 1600 11.62 Pliensbachian Statherian Lower 407.6 ±2.6 Serravallian 13.82 190.8 ±1.0 Lower 1800 Miocene Pragian 410.8 ±2.8 Langhian Sinemurian Proterozoic Neogene 15.97 Orosirian 199.3 ±0.3 Lochkovian Paleo- Hettangian 2050 Burdigalian 201.3 ±0.2 419.2 ±3.2 proterozoic 20.44 Mesozoic Rhaetian Pridoli Rhyacian Aquitanian 423.0 ±2.3 23.03 ~ 208.5 Ludfordian 2300 Cenozoic Chattian Ludlow 425.6 ±0.9 Siderian 28.1 Gorstian Oligocene Upper Norian 427.4 ±0.5 2500 Rupelian Wenlock Homerian
  • The Challenge of the Aeronian/Telychian (Llandovery) Boundary

    The Challenge of the Aeronian/Telychian (Llandovery) Boundary

    GEOLOGICA BALCANICA, 46 (2), Sofia, Nov. 2017, pp. 3–10. The Silurian stage boundaries in Bulgaria: the challenge of the Aeronian/Telychian (Llandovery) boundary Valeri Sachanski University of Mining and Geology, Department of Geology and Geoinformatics, Studentski Grad, Prof. Boyan Kamenov Str., 1700 Sofia, Bulgaria; Geological Institute, Bulgarian Academy of Sciences, Acad. G. Bonchev Str., Bl. 24, 1113 Sofia, Bulgaria; e-mail: [email protected] (Accepted in revised form: October 2017) Abstract. The upper Aeronian to lower Telychian (Llandovery Series, lower Silurian) strata in the paras- tratotype for the Saltar Formation (chert-shale succession) are documented in terms of their lithologies and graptolite biostratigraphy. The studied section Asaritsa сrops out in the western sector of the Stara Planina Mountains (Svoge Unit of the Srednogorie Zone). In section Asaritsa, the highest Aeronian graptolitic strata are separated from the lowest Telychian graptolitic strata by a graptolite-barren interval of pale-coloured shale. Based on the graptolites documented below and above this interval, as well as on its correlation to sec- tions studied in detail from the Czech Republic and Spain, the immediate underlay of the pale-coloured shale is referred to the uppermost part of the sedgwickii Zone, the complete pale-coloured shale interval, or its great- est part, to the lower guerichi Zone, and the directly overlying it succession to the upper part of the guerichi Zone. The Aeronian/Telychian boundary is assumed to lie at the base of this pale-coloured shale package, or within its lower part. The study provides documentation of Metaclimacograptus undulatus, Parapetalolithus praecedens, Rastrites carnicus, Rivagraptus bellulus, and illustrations of Stimulograptus for the first time in Bulgaria.

  • Paleogeographic Maps Earth History

    History of the Earth Age AGE Eon Era Period Period Epoch Stage Paleogeographic Maps Earth History (Ma) Era (Ma) Holocene Neogene Quaternary* Pleistocene Calabrian/Gelasian Piacenzian 2.6 Cenozoic Pliocene Zanclean Paleogene Messinian 5.3 L Tortonian 100 Cretaceous Serravallian Miocene M Langhian E Burdigalian Jurassic Neogene Aquitanian 200 23 L Chattian Triassic Oligocene E Rupelian Permian 34 Early Neogene 300 L Priabonian Bartonian Carboniferous Cenozoic M Eocene Lutetian 400 Phanerozoic Devonian E Ypresian Silurian Paleogene L Thanetian 56 PaleozoicOrdovician Mesozoic Paleocene M Selandian 500 E Danian Cambrian 66 Maastrichtian Ediacaran 600 Campanian Late Santonian 700 Coniacian Turonian Cenomanian Late Cretaceous 100 800 Cryogenian Albian 900 Neoproterozoic Tonian Cretaceous Aptian Early 1000 Barremian Hauterivian Valanginian 1100 Stenian Berriasian 146 Tithonian Early Cretaceous 1200 Late Kimmeridgian Oxfordian 161 Callovian Mesozoic 1300 Ectasian Bathonian Middle Bajocian Aalenian 176 1400 Toarcian Jurassic Mesoproterozoic Early Pliensbachian 1500 Sinemurian Hettangian Calymmian 200 Rhaetian 1600 Proterozoic Norian Late 1700 Statherian Carnian 228 1800 Ladinian Late Triassic Triassic Middle Anisian 1900 245 Olenekian Orosirian Early Induan Changhsingian 251 2000 Lopingian Wuchiapingian 260 Capitanian Guadalupian Wordian/Roadian 2100 271 Kungurian Paleoproterozoic Rhyacian Artinskian 2200 Permian Cisuralian Sakmarian Middle Permian 2300 Asselian 299 Late Gzhelian Kasimovian 2400 Siderian Middle Moscovian Penn- sylvanian Early Bashkirian
  • 2009 Geologic Time Scale Cenozoic Mesozoic Paleozoic Precambrian Magnetic Magnetic Bdy

    2009 Geologic Time Scale Cenozoic Mesozoic Paleozoic Precambrian Magnetic Magnetic Bdy

    2009 GEOLOGIC TIME SCALE CENOZOIC MESOZOIC PALEOZOIC PRECAMBRIAN MAGNETIC MAGNETIC BDY. AGE POLARITY PICKS AGE POLARITY PICKS AGE PICKS AGE . N PERIOD EPOCH AGE PERIOD EPOCH AGE PERIOD EPOCH AGE EON ERA PERIOD AGES (Ma) (Ma) (Ma) (Ma) (Ma) (Ma) (Ma) HIST. HIST. ANOM. ANOM. (Ma) CHRON. CHRO HOLOCENE 65.5 1 C1 QUATER- 0.01 30 C30 542 CALABRIAN MAASTRICHTIAN NARY PLEISTOCENE 1.8 31 C31 251 2 C2 GELASIAN 70 CHANGHSINGIAN EDIACARAN 2.6 70.6 254 2A PIACENZIAN 32 C32 L 630 C2A 3.6 WUCHIAPINGIAN PLIOCENE 260 260 3 ZANCLEAN 33 CAMPANIAN CAPITANIAN 5 C3 5.3 266 750 NEOPRO- CRYOGENIAN 80 C33 M WORDIAN MESSINIAN LATE 268 TEROZOIC 3A C3A 83.5 ROADIAN 7.2 SANTONIAN 271 85.8 KUNGURIAN 850 4 276 C4 CONIACIAN 280 4A 89.3 ARTINSKIAN TONIAN C4A L TORTONIAN 90 284 TURONIAN PERMIAN 10 5 93.5 E 1000 1000 C5 SAKMARIAN 11.6 CENOMANIAN 297 99.6 ASSELIAN STENIAN SERRAVALLIAN 34 C34 299.0 5A 100 300 GZELIAN C5A 13.8 M KASIMOVIAN 304 1200 PENNSYL- 306 1250 15 5B LANGHIAN ALBIAN MOSCOVIAN MESOPRO- C5B VANIAN 312 ECTASIAN 5C 16.0 110 BASHKIRIAN TEROZOIC C5C 112 5D C5D MIOCENE 320 318 1400 5E C5E NEOGENE BURDIGALIAN SERPUKHOVIAN 326 6 C6 APTIAN 20 120 1500 CALYMMIAN E 20.4 6A C6A EARLY MISSIS- M0r 125 VISEAN 1600 6B C6B AQUITANIAN M1 340 SIPPIAN M3 BARREMIAN C6C 23.0 345 6C CRETACEOUS 130 M5 130 STATHERIAN CARBONIFEROUS TOURNAISIAN 7 C7 HAUTERIVIAN 1750 25 7A M10 C7A 136 359 8 C8 L CHATTIAN M12 VALANGINIAN 360 L 1800 140 M14 140 9 C9 M16 FAMENNIAN BERRIASIAN M18 PROTEROZOIC OROSIRIAN 10 C10 28.4 145.5 M20 2000 30 11 C11 TITHONIAN 374 PALEOPRO- 150 M22 2050 12 E RUPELIAN
  • Integrated Biostratigraphy of the Lower Silurian of the Aizpute-41 Core, Latvia

    Integrated Biostratigraphy of the Lower Silurian of the Aizpute-41 Core, Latvia

    Geol. Mag. 140 (2), 2003, pp. 205–229. c 2003 Cambridge University Press 205 DOI: 10.1017/S0016756802007264 Printed in the United Kingdom Integrated biostratigraphy of the lower Silurian of the Aizpute-41 core, Latvia D. K. LOYDELL*, P. MANNIK¨ † &V.NESTOR† *School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Road, Portsmouth PO1 3QL, UK †Institute of Geology, Tallinn Technical University, 7 Estonia Avenue, 10143 Tallinn, Estonia (Received 20 November 2001; accepted 28 November 2002) Abstract –Integrated graptolite, conodont and chitinozoan biostratigraphical data is presented from the Rhuddanian through to lower Sheinwoodian of the Aizpute-41 core, Latvia. Correlation of the biozonation schemes based upon the three groups is achieved from the cyphus through to lowermost riccartonensis graptolite biozones, except for the upper Aeronian and lower Telychian, which lack both chitinozoans and graptolites, and upper lapworthi through to approximately base murchisoni graptolite Biozone, where there is interpreted to be an unconformity. Datum 2 of the Ireviken Event is correlated with a level at the base of or within the murchisoni Biozone. It is possible that the changes in conodont assemblages at Datum 2 on Gotland are the result of an unconformity here. Streptograptus? kaljoi sp. nov., from the lower spiralis graptolite Biozone, is described. Keywords: graptolites, Chitinozoa, Conodonta, Silurian, biostratigraphy. 1. Introduction setting is further off-shore than that of Ohesaare (Fig. 1; see Loydell, Kaljo & Mannik,¨ 1998, for details of the Recognition of the importance of integrated biostrati- Ohesaare core), and the succession within the core graphical studies in the precise cross-facies correlation is significantly more complete, most notably in the required to test models of biotic and facies change in Aeronian.
  • Rendspi 1-Studies

    Rendspi 1-Studies

    Rendiconti della Società Paleontologica Italiana 3 (I) The Silurian of Sardinia volume in honour of Enrico Serpagli Edited by Carlo Corradini Annalisa Ferretti Petr Storch Società Paleontologica Italiana 2009 I Rendiconti della Società Paleontologica Italiana, 3 (1), 2009: 43-49 The Silurian of the External Nappes (southeastern Sardinia) CARLO CORRADINI, ANNALISA FERRETTI C. Corradini - Dipartimento di Scienze della Terra, Università di Cagliari, via Trentino 51, I-09127 Cagliari (Italy); [email protected] A. Ferretti - Dipartimento di Scienze della Terra, Università di Modena e Reggio Emilia, largo S. Eufemia 19, I-41100 Modena (Italy);[email protected] ABSTRACT - The most complete and best known Silurian succession of southeastern Sardinia is exposed in the Gerrei tectonic Unit. The Silurian sequence starts with Rhuddanian-lowermost Gorstian black graptolitic shales (Lower Graptolitic Shales), followed by a lower Gorstian- end Pridoli nodular calcareous unit (Ockerkalk). Graptolitic shales (Upper Graptolitic Shales) document the Lower Devonian. An integrated scheme of graptolite and conodont biozonation, compiled from Sardic data, is provided. KEY WORDS - Silurian, southeastern Sardinia, biostratigraphy, Gerrei tectonic Unit, Lower Graptolitic Shales, Ockerkalk. INTRODUCTION Silurian rocks are quite widespread in southeastern Sardinia, where they crop out widely in the Gerrei tectonic Unit and more rarely in the other tectonic units of the area (Sarrabus and Meana Sardo tectonic units). In the first approximation, the sequence is constituted by black graptolitic shales in the lower part, followed by nodular limestones of late Silurian age. THE GERREI TECTONIC UNIT The Gerrei tectonic Unit spans from middle Cambrian to Lower Carboniferous and includes the most complete middle Palaeozoic sequence of Sardinia (Fig.
  • Late Silurian Trilobite Palaeobiology And

    Late Silurian Trilobite Palaeobiology And

    LATE SILURIAN TRILOBITE PALAEOBIOLOGY AND BIODIVERSITY by ANDREW JAMES STOREY A thesis submitted to the University of Birmingham for the degree of DOCTOR OF PHILOSOPHY School of Geography, Earth and Environmental Sciences University of Birmingham February 2012 University of Birmingham Research Archive e-theses repository This unpublished thesis/dissertation is copyright of the author and/or third parties. The intellectual property rights of the author or third parties in respect of this work are as defined by The Copyright Designs and Patents Act 1988 or as modified by any successor legislation. Any use made of information contained in this thesis/dissertation must be in accordance with that legislation and must be properly acknowledged. Further distribution or reproduction in any format is prohibited without the permission of the copyright holder. ABSTRACT Trilobites from the Ludlow and Přídolí of England and Wales are described. A total of 15 families; 36 genera and 53 species are documented herein, including a new genus and seventeen new species; fourteen of which remain under open nomenclature. Most of the trilobites in the British late Silurian are restricted to the shelf, and predominantly occur in the Elton, Bringewood, Leintwardine, and Whitcliffe groups of Wales and the Welsh Borderland. The Elton to Whitcliffe groups represent a shallowing upwards sequence overall; each is characterised by a distinct lithofacies and fauna. The trilobites and brachiopods of the Coldwell Formation of the Lake District Basin are documented, and are comparable with faunas in the Swedish Colonus Shale and the Mottled Mudstones of North Wales. Ludlow trilobite associations, containing commonly co-occurring trilobite taxa, are defined for each palaeoenvironment.