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The giants of the phylum Brachiopoda: a matter of diet?

Angiolini, Lucia; Crippa, Gaia; Azmy, Karem; Capitani, Giancarlo; Confalonieri, Giorgia; Della Porta, Giovanna; Griesshaber, Erika; Harper, David A.T.; Leng, Melanie J.; Nolan, Leah; Orlandi, Marco; Posenato, Renato; Schmahl, Wolfgang W.; Banks, Vanessa J.; Stephenson, Michael H.

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Lucia Angiolini

Gaia Crippa

Karem Azmy

Giancarlo Capitani

Giorgia Confalonieri

Giovanna Della Porta

Erika Griesshaber

Melanie J. Leng

Leah Nolan

Marco Orlandi

Renato Posenato

Wolfgang W. Schmahl

Vanessa J. Banks

Michael H. Stephenson


The species of the brachiopod Gigantoproductus are giants within the Palaeozoic sedentary benthos. This presents a dilemma as living brachiopods have low‐energy lifestyles. Although brachiopod metabolic rates were probably higher during the Palaeozoic than today, the massive size reached by species of Gigantoproductus is nevertheless unusual. By examining the diet of Gigantoproductus species from the Visean (Mississippian, Carboniferous) of Derbyshire (UK), we seek to understand the mechanisms that enabled those low‐metabolism brachiopod species to become giants. Were they suspension feeders, similar to all other brachiopods, or did endosymbiosis provide a lifestyle that allowed them to have higher metabolic rates and become giants? We suggest that the answer to this conundrum may be solved by the identification of the biogeochemical signatures of symbionts, through combined analyses of the carbon and nitrogen‐isotopic compositions of the occluded organic matrix within their calcite shells. The shells are formed of substructured columnar units that are remarkably long and a few hundreds of microns wide, deemed to be mostly pristine based on multiple analyses (petrography, cathodoluminescence (CL), scanning electron microscopy (SEM), electron backscatter diffraction (EBSD), transmission electron microscopy (TEM)); they contain occluded organic fractions detected by TEM, nuclear magnetic resonance (NMR) and gas chromatography mass spectrometry (GC‐MS) analyses. We conclude that the gigantic size reached by the species of Gigantoproductus is probably the result of a mixotroph lifestyle, by which they could rely on the energy and nutrients derived both from photosymbiotic microbes and from filtered particulate food.

Journal Article Type Article
Acceptance Date Feb 19, 2019
Online Publication Date Jun 10, 2019
Publication Date 2019-11
Deposit Date Jun 24, 2019
Publicly Available Date Jun 10, 2020
Journal Palaeontology
Print ISSN 0031-0239
Electronic ISSN 1475-4983
Publisher Wiley
Peer Reviewed Peer Reviewed
Volume 62
Issue 6
Pages 889-917
Public URL


Accepted Journal Article (7.8 Mb)

Copyright Statement
This is the peer reviewed version of the following article: Angiolini, Lucia, Crippa, Gaia, Azmy, Karem, Capitani, Giancarlo, Confalonieri, Giorgia, Della Porta, Giovanna, Griesshaber, Erika, Harper, David A. T., Leng, Melanie J., Nolan, Leah, Orlandi, Marco, Posenato, Renato, Schmahl, Wolfgang W., Banks, Vanessa J. & Stephenson, Michael H. (2019). The giants of the phylum Brachiopoda: a matter of diet? Palaeontology 62(6): 889-917 which has been published in final form at This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions.

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