Permophiles Issue #66 Supplement 1 pods which may further address these questions. Glenn A. Brock The world’s oceans are changing. IPCC (2013) predictions Department of Biological Sciences, Macquarie University, NSW suggest that by the end of the current century our seas will be 2109, Australia ZDUPHUDQGWKHLUS+VLJQL¿FDQWO\ORZHU$OWKRXJKWKLVZLOOEH a challenge to all organisms, how will it impact brachiopods? Leonid E. Popov Given that they have a higher proportion of mineralised tissue 'HSDUWPHQWRI*HRORJ\1DWLRQDO0XVHXPRI:DOHV&DUGL൵ than virtually any other invertebrate group what will be the CF10 3NP, UK WKUHDWWRWKHPHLWKHULQWHUPVRIJURZLQJWKHLUVKHOOVLQWKH¿UVW Brachiopods are richly represented in the rock record and instance or repairing and maintaining that shell once it is made? as early as the Cambrian, where they show an impressive diver- In this talk I will review a series of experiments and historical sity of form and in shell morphology (e.g., Harper et al., 2017). studies undertaken with Emma Cross and Lloyd Peck (Cross et 3UHVHQWO\ WKH JURXS LV ¿UPO\ URRWHG ZLWKLQ WKH ORSKRWURFKR- al., 2015, 2016, 2018) that seek to explore the answers to these zoan branch of the bilaterian tree based on molecular data. Our questions. UHVHDUFKKDVLGHQWL¿HGVRPHPHPEHUVRIWKHHQLJPDWLF(DUO\ References Cambrian organophosphatic tommotiids as belonging to the &URVV(/3HFN/6 +DUSHU(02FHDQDFLGL¿FD- brachiopod stem (e.g., Holmer et al., 2002). Subsequent discov- tion does not impact shell growth or repair of the Antarctic HULHVRIWKH¿UVWHYHUDUWLFXODWHGVFOHULWRPHVRIEccentrotheca, brachiopod Liothyrella uva (Broderip, 1833). Journal of Paterimitra, and the inferred bivalved scleritome of Micrina f rom Experimental Marine Biology and Ecology, 462, 29–35. the lower Cambrian of South Australia reveals these three tom- Cross, E.L., Peck, L.S., Lamare, M.D. & Harper, E.M. 2016. PRWLLGWD[DDVOLNHO\ORSKRSKRUDWHVHVVLOH¿OWHUIHHGHUVZKLFK 1RRFHDQDFLGL¿FDWLRQH൵HFWVRQVKHOOJURZWKDQGUHSDLULQ in the Paterimitra includes a preserved pedicle tube (Skovsted the New Zealand brachiopod Calloria inconspicua (Sowerby, et al., 2008, 2009; Holmer et al., 2008, 2011). It is likely that the 1846). ICES Journal of Marine Sciences, 73, 920-926. Brachiopoda may have evolved, by sclerite reduction and modi- Cross, E. L., Harper, E. M. & Peck, L. S. 2018. A 120-year ¿FDWLRQIURPDVHVVLOHVFOHULWRPHWXEHGZHOOLQJEccentrotheca. record of resilience to environmental change in brachio- Micrina and Paterimitra also preserve traces of their earliest pods. Global Change Biology. ISSN 1354-1013 DOI 10.1111/ ontogeny, including bivalved larval shells with evidence of the gcb.14085 earliest larval attachment. The conjoined bivalved shell of adult Harper, E.M. & Peck, L.S. 2016. Latitudinal and depth gra- living brachiopods most likely represents a plesiomorphic char- dients in marine predation pressure. Global Ecology and acter retained from planktic tommotiid larvae; the crown group Biogeography 25, 670-678. body plan probably evolved independently in living linguliform IPCC. 2013. Climate change 2013: The physical science basis. In and rhynchonelliforms (Holmer et al., 2011; see also Zhang et T. F. Stocker, D. Qin, G-K. Plattner, M. Tignor, S. K. Allen, J. al., 2018a,b). Studies of new records of exceptionally preserved Boschung, A. Nauels, Y. Xia, V. Bex & P. M. Midgley (Eds.), soft-bodied fossil assemblages (Konservat-Lagerstätten) are :RUNLQJJURXS,FRQWULEXWLRQWRWKH¿IWKDVVHVVPHQWUHSRUWRI also critical. For example, we have now been able to reassess the intergovernmental panel on climate change (pp. 1–1522). highly problematic Chengjiang fossils such as the Cotyledion Cambridge, United Kingdom and New York, NY, USA: as a sclerite-covered sessile tube-shaped probable stem Cambridge University Press. “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 Plenary lecture to our understanding of more derived brachiopods, such as lin- Brachiopod Phylogeny in the Cambrian guliforms, obolellates and rhynchonelliforms (e.g., Zhang et al., 2009, 2014, 2015; Holmer et al., 2018a). Exceptionally pre- Lars E. Holmer served pedunculated Cambrian rhynchonelliforms from both Shaanxi Key laboratory of Early Life and Environments and the Chengjiang and Utah also indicate that early members of Department of Geology, State Key Laboratory of Continental this most important group of living brachiopods had two widely Dynamics, Northwest University, Xi’an 710069, China GL൵HUHQWW\SHVRISHGLFOHV +ROPHUHWDODE 'HVSLWH Department of Earth Sciences, Palaeobiology, Uppsala WKHVHQHZ¿QGLQJVPDQ\DVSHFWVRIWKHSK\ORJHQ\DQGUDGLD- University, SE-752 36 Uppsala, Sweden tion of the Brachiopoda in the Cambrian remain problematic and controversial issues. Presently only two of the three major bra- Zhifei Zhang chiopod clades, the Linguliformea and the Rhynchonelliformea, Shaanxi Key laboratory of Early Life and Environments and have known Cambrian representatives. The origin and phylog- Department of Geology, State Key Laboratory of Continental eny of the Craniiformea remains problematic, but it is possible Dynamics, Northwest University, Xi’an 710069, China that Cambrian rhynchonelliform chileate-like brachiopods are involved in the processes. Zhiliang Zhang Shaanxi Key laboratory of Early Life and Environments and References Department of Geology, State Key Laboratory of Continental Harper, D., Popov, L. & Holmer, L.E. 2017. Brachiopods, origin Dynamics, Northwest University, Xi’an 710069, China 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¿FDWLRQSDWWHUQVRIEUDFKLRSRGV group brachiopod from the Lower Cambrian – support for a Micrina (halkieriid) ancestry. Palaeontology, 45, 875-882. after the end Ordovician mass extinction Holmer, L. E., Skovsted, C. B., Brock, G. A., Valentine, J.L. & DQGLWVSDODHRELRJHRJUDSKLFVLJQL¿FDQFH Paterson, J. R. 2008. The Early Cambrian tommotiid Micrina, a sessile bivalved stem group brachiopod. Biological letters, Bing Huang 4, 724-728. State Key Laboratory of Paleobiology and Stratigraphy, Nanjing Holmer, L.E., Skovsted, C.B., Larsson, C., Brock, G.A. & Institute of Geology and Palaeontology, The Chinese Academy Zhang, Z. 2011. First record of a bivalve larval shell in Early of Sciences, Nanjing 210008, PR China &DPEULDQ WRPPRWLLGV DQG LWV SK\ORJHQHWLF VLJQL¿FDQFH Jisuo Jin Palaeontology, 54, 235-239. 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 Jiayu Rong WKHLUSK\ORJHQHWLFVLJQL¿FDQFHJournal of Paleontology (in State Key Laboratory of Paleobiology and Stratigraphy, Nanjing press). Institute of Geology and Palaeontology, The Chinese Academy Holmer, L.E., Popov, L., Ghobadi Pour, M., Claybourn, T., of Sciences, Nanjing 210008, PR China Zhang, Z., Brock, G.A. & Zhang, Z. 2018b. Evolutionary sig- QL¿FDQFHRIDPLGGOH&DPEULDQ 6HULHV LQVLWXRFFXUUHQFH The Late Ordovician biotic crisis was associated with a brief of the pedunculate rhynchonelliform brachiopod Nisusia sul- but intense glaciation episode in earth history. Post-glacial marine cata. Lethaia (in press). transgression created vast habits in epicontinental seas, in which Skovsted, C, Brock, G, Paterson, J, Holmer, L. & Budd, G 2008 EHQWKLFIDXQDVHVWDEOLVKHGDQGGLYHUVL¿HGZLWKEUDFKLRSRGVEHLQJ The scleritome of Eccentrotheca from the Lower Cambrian one of the most abundant and diverse fossil groups. Based on RI6RXWK$XVWUDOLDORSKRSKRUDWHD൶QLWLHVDQGLPSOLFDWLRQV detailed data of brachiopod occurrences after the end-Ordovician for tommotiid phylogeny. Geology, 36, 171-174. mass extinction, together with newly published data, we analyzed Skovsted, C.B., Holmer, L.E., Larsson, C.M, Högström, A.E.S., WKHGLYHUVL¿FDWLRQSDWWHUQVRIEUDFKLRSRGVIURPWKH5KXGGDQLDQ Brock, G.A., Topper, T.P., Balthasar, U., Petterson Stolk, S. to the Aeronian (early Silurian) by Network Analysis (Fig. 1) and & Paterson, J.R.. 2009. The scleritome of Paterimitra: an Frequency Distribution Analysis. Rhuddanian brachiopod faunas early Cambrian stem group brachiopod from South Australia. were characterized by low diversity and localized high abundance, Proceedings of the Royal Society B, 276, 1651-1656. except for a relatively high-diversity early Rhuddanian fauna in Zhang, Z.F., Li, G.X., Emig, C.C., Han, J., Holmer, L.E. & Shu, Avalonia-Baltica and South China. Invariably, these faunas were D.G. 2009. Architecture and function of the lophophore predominated by re-established holdover and once-cosmopolitan in the problematic brachiopod Heliomedusa orienta (Early taxa from the Late Ordovician, primarily orthides and stropho- Cambrian, South China). Geobios, 42: 649-661. menides. By the Aeronian, global brachiopod diversity nearly Zhang, Z., Holmer, L.E., Skovsted, C., Brock, G. & Budd, G. doubled in comparison with the Rhuddanian, owing to a major 2013. A sclerite-bearing stem group entoproct from the early GLYHUVL¿FDWLRQ RI 6LOXULDQW\SH EUDFKLRSRGV LQ ERWK WKH SDOHR- Cambrian and its implications. 6FLHQWL¿F5HSRUWV 3, 1066. tropics and high-latitude Gondwana, as well as the concomitant Zhang, Z.-F., Li, G.-X., Holmer, L.E., Brock, G.A., Balthasar, U., proliferation of endemic and cosmopolitan taxa, in association Skovsted, C.B., Fu, D-J., Zhang, X.-L., Wang, H.-Z., Butler, with a global expansion of epicontinental seas and heterogeneity A., Zhang, Z.-L., Cao, C-Q., Han, J., Liu, J.-N. & Shu, D.-G. of specialized local habitats. A drastic turnover of brachiopods 2014. An early Cambrian agglutinated tubular lophophorate IURPWKH2UGRYLFLDQWRWKH6LOXULDQW\SHVLJQL¿FDQWO\D൵HFWHGWKH with brachiopod characters. 6FLHQWL¿F5HSRUWV, 4682. Paleozoic evolutionary fauna. With the ordinal data of brachio- Zhang Z.F., Zhang, Z.L., Holmer , L.E. & Li, G.X. 2015. First pods in major palaeoplates, we discussed the turnover process. report of linguloid brachiopods with soft parts from the lower Compared with Ordovician orthides and strophomenides, which Cambrian (Series 2, Stage 4) of the Three Gorges area, South globally increased their diversity after the extinction, atrypides China. Annales de Paléontologie, 101:167–177. and pentamerides of Silurian group displayed a drastic diversi- Zhang, Z., Zhang, Z., Holmer, L.E. & Chen, F. 2018a. Post- ¿FDWLRQ IURP 5KXGGDQLDQ WR $HURQLDQ RQO\ LQ ORZHU ODWLWXGH metamorphic allometry in the earliest acrotretoid brachiopods continents, demonstrating the two new orders preferred more from the lower Cambrian (Series 2) of South China, and its warm water environments than the Ordovician groups. Atrypides implications. Palaeontology, 61, 183–207. UHFRYHUHGIURPWKHHDUO\5KXGGDQLDQZKHUHDVGLYHUVL¿FDWLRQRI Zhang, Z., Popov, L., Holmer, L.E. & Zhang, Z. 2018b. Earliest pentamerides delayed until to Aeronian, and this is typically dem- RQWRJHQ\RIHDUO\&DPEULDQDFURWUHWRLGEUDFKLRSRGV±¿UVW onstrated by the record of many endemic taxa from South China. evidence for metamorphosis and its implications. BMC evo- The Silurian-type atrypide and pentameride communities thrived lutionary biology (in press). 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).

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