<documentid="958002F20F64EA8E0DA9A7E2134F7C44"ID-CLB-Dataset="305524"ID-DOI="10.5852/cr-palevol2024v23a22"ID-GBIF-Dataset="fefdf6a6-89e6-4b61-8d87-ff08580d85f9"ID-ISSN="1777-571X"ID-Zenodo-Dep="13595789"ID-ZooBank="urn:lsid:zoobank.org:pub:FAE07554-6F51-46B5-A81D-EA7D9091E776"IM.bibliography_approvedBy="valdenar"IM.illustrations_approvedBy="felipe"IM.materialsCitations_approvedBy="felipe"IM.metadata_approvedBy="felipe"IM.taxonomicNames_approvedBy="valdenar"IM.treatments_approvedBy="valdenar"checkinTime="1732638299342"checkinUser="felipe"docAuthor="Uchman, Alfred & Wetzel, Andreas"docDate="2024"docId="03DB87D8FFEEFF90ACF5FB19027DC333"docLanguage="en"docName="CRPalevol.23.22.325-338.pdf"docOrigin="Comptes Rendus Palevol 23 (22)"docSource="http://dx.doi.org/10.5852/cr-palevol2024v23a22"docStyle="DocumentStyle:DACF86F01658A8850E13A63D98C287FD.1:CRPalevol.2020-.journal_article"docStyleId="DACF86F01658A8850E13A63D98C287FD"docStyleName="CRPalevol.2020-.journal_article"docStyleVersion="1"docTitle="Zoophycos Massalongo 1855"docType="treatment"docVersion="5"lastPageNumber="328"masterDocId="FFE2FFA0FFEBFF95AC1CFF8D064AC71C"masterDocTitle="Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows"masterLastPageNumber="338"masterPageNumber="325"pageNumber="328"updateTime="1734625551218"updateUser="ExternalLinkService"zenodo-license-document="CC-BY-4.0">
<mods:titleid="447C06BBD85C38C015599F3AF3DC205D">Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows</mods:title>
<mods:affiliationid="F6D036B55FE685B20D5E61EE3190A222">Jagiellonian University, Faculty of Geography and Geology, Institute of Geological Sciences, Gronostajowa 3 a, PL- 30 - 387 Kraków (Poland) alfred. uchman @ uj. edu. pl (corresponding author)</mods:affiliation>
<bibRefCitationid="EFE34B3FFFEEFF90ADC0FB1904E8C3B3"author="MASSALONGO A. D. B."box="[476,674,1172,1199]"pageId="5"pageNumber="328"refId="ref11232"refString="MASSALONGO A. D. B. 1855. - Zoophycos, novum genus plantarum fossilium. Antonelli, Verona, 52 p."type="book"year="1855">Massalongo, 1855</bibRefCitation>
<figureCitationid="13492A4BFFEEFF90ADBAFB3907AFC3D2"box="[422,485,1204,1230]"captionStart="FIG"captionStartId="6.[132,143,1656,1673]"captionTargetBox="[132,1455,215,1609]"captionTargetId="figure-311@6.[603,1266,771,960]"captionTargetPageId="6"captionText="FIG. 1. — Sequestered and downward-transferred material in Zoophycos Massalongo,1855: A, black mud and white nannofossil ooze displaced for> 26 (black arrows) and> 23 cm (white arrows) downward, respectively, as indicated by colour and composition of sediment (Institute of Geological Sciences Kiel, Germany, core 16867-3; 2.2°S, 5.1°E, 3894 m water depth, mid Atlantic; 16.56-16.99 m core depth); B, sequestered organic-rich grey sediment filling Zoophycos spreite (white arrows) produced in red pelagic mud (Maastrichtian, Zumaia, Spain); C, homogeneous and pelleted sequestered sediment alternating with host sediment in Zoophycos spreite; inset (C’) showing pellets occurring in both sequestered sediment (green arrows) and host sediment (yellow arrows) (DSDP Leg 93, Site 605, Core 16-4, 59.5-66.5 cm core depth; for details see Wetzel 1987); D, truncation of lamellae (marked with red stippled lines) in Zoophycos spreite in horizontal section, indicating reworking of previously emplaced material (Paleogene, Zumaia, Spain). Scale bars: 1 cm."figureDoi="http://doi.org/10.5281/zenodo.14530602"httpUri="https://zenodo.org/record/14530602/files/figure.png"pageId="5"pageNumber="328">Fig. 1</figureCitation>
represents a regularly to irregularly coiled, simple to complex lobate spreite burrow that has shown changes in shape, size, and geometry through the Phanerozoic (e.g.
<bibRefCitationid="EFE34B3FFFEEFF90AEB5FAB90680C272"author="SEILACHER A."pageId="5"pageNumber="328"pagination="359 - 376"refId="ref12450"refString="SEILACHER A. 1977 b. - Evolution of trace fossil communities, in HALLAM A. (ed.), Patterns of evolution as illustrated by the fossil record. Developments in Palaeontology and Stratigraphy 5: 359 - 376. https: // doi. org / 10.1016 / S 0920 - 5446 (08) 70331 - 5"type="journal article"year="1977">Seilacher 1977b</bibRefCitation>
; later refined by
<bibRefCitationid="EFE34B3FFFEEFF90AD9EFAD90405C272"author="CHAMBERLAIN C. K."box="[386,591,1364,1390]"pageId="5"pageNumber="328"pagination="13 - 22"refId="ref8804"refString="CHAMBERLAIN C. K. 2000. - Prologue to the study of Zoophycos. Ichnology Newsletter 22: 13 - 22."type="journal article"year="2000">Chamberlain 2000</bibRefCitation>
and
<bibRefCitationid="EFE34B3FFFEEFF90AE9AFAD906F7C292"author="ZHANG L. - J. & FAN R. - Y. & GONG Y. - M."pageId="5"pageNumber="328"refId="ref14979"refString="ZHANG L. - J., FAN R. - Y. & GONG Y. - M. 2015. - Zoophycos macroevolution since 541 Ma. Scientific Reports 5: 14954. https: // doi. org / 10.1038 / srep 14954"type="journal volume"year="2015">
older than the late Mesozoic, different ways of spreite production and behaviour are possible, including fodinichnial behaviour (e.g.
<bibRefCitationid="EFE34B3FFFEEFF90AE28FA3E06F7C2F1"author="OLIVERO D. & GAILLARD C."pageId="5"pageNumber="328"pagination="249 - 260"refId="ref12035"refString="OLIVERO D. & GAILLARD C. 1996. - Paleoecology of Jurassic Zoophycos from south-eastern France. Ichnos 4 (4): 249 - 260. https: // doi. org / 10.1080 / 10420949609380135"type="journal article"year="1996">Olivero & Gaillard 1996</bibRefCitation>
Evidence for sequestration of sediment on the seafloor is given by the colour of the spreite fill in combination with enlarged organic carbon values. The spreite consists of alternating lamellae containing host sediment and material likely sequestered on the seafloor and transferred downward; pellets may occur in both
<figureCitationid="13492A4BFFEEFF90AE17F95F0402C1F1"box="[523,584,1746,1773]"captionStart="FIG"captionStartId="6.[132,143,1656,1673]"captionTargetBox="[132,1455,215,1609]"captionTargetId="figure-311@6.[603,1266,771,960]"captionTargetPageId="6"captionText="FIG. 1. — Sequestered and downward-transferred material in Zoophycos Massalongo,1855: A, black mud and white nannofossil ooze displaced for> 26 (black arrows) and> 23 cm (white arrows) downward, respectively, as indicated by colour and composition of sediment (Institute of Geological Sciences Kiel, Germany, core 16867-3; 2.2°S, 5.1°E, 3894 m water depth, mid Atlantic; 16.56-16.99 m core depth); B, sequestered organic-rich grey sediment filling Zoophycos spreite (white arrows) produced in red pelagic mud (Maastrichtian, Zumaia, Spain); C, homogeneous and pelleted sequestered sediment alternating with host sediment in Zoophycos spreite; inset (C’) showing pellets occurring in both sequestered sediment (green arrows) and host sediment (yellow arrows) (DSDP Leg 93, Site 605, Core 16-4, 59.5-66.5 cm core depth; for details see Wetzel 1987); D, truncation of lamellae (marked with red stippled lines) in Zoophycos spreite in horizontal section, indicating reworking of previously emplaced material (Paleogene, Zumaia, Spain). Scale bars: 1 cm."figureDoi="http://doi.org/10.5281/zenodo.14530602"httpUri="https://zenodo.org/record/14530602/files/figure.png"pageId="5"pageNumber="328">Fig. 1</figureCitation>
). The sequestered sediment is commonly darker in greenish-grey host sediment or grey in reddish host sediment. In both cases, the organic carbon content of the spreite fill is higher than that of the host sediment; for example, the black spreite material of Paleocene
<bibRefCitationid="EFE34B3FFFEEFF90ADE6F83F04B4C0D0"author="WETZEL A. & UCHMAN A."box="[506,766,1970,1996]"pageId="5"pageNumber="328"pagination="351 - 369"refId="ref14463"refString="WETZEL A. & UCHMAN A. 1998. - Biogenic sedimentary structures in mudstones - an overview, in SCHIEBER J., ZIMMERLE W. & SETHI P. (eds), Shales and Mudstones I. Schweizerbart, Stuttgart: 351 - 369."type="book chapter"year="1998">Wetzel & Uchman 1998</bibRefCitation>
: fig. 1d, e). In the Eocene of Arnakatxa Headland near Bilbao, 0.3-0.7%
in red limestone lutite alternations. However, in deep oceanic settings, also lighter material deposited during periods of enhanced nannoplankton productivity can be transferred downward (
<figureCitationid="13492A4BFFEEFF90AFAEFEDA0243C66D"box="[946,1033,343,369]"captionStart="FIG"captionStartId="6.[132,143,1656,1673]"captionTargetBox="[132,1455,215,1609]"captionTargetId="figure-311@6.[603,1266,771,960]"captionTargetPageId="6"captionText="FIG. 1. — Sequestered and downward-transferred material in Zoophycos Massalongo,1855: A, black mud and white nannofossil ooze displaced for> 26 (black arrows) and> 23 cm (white arrows) downward, respectively, as indicated by colour and composition of sediment (Institute of Geological Sciences Kiel, Germany, core 16867-3; 2.2°S, 5.1°E, 3894 m water depth, mid Atlantic; 16.56-16.99 m core depth); B, sequestered organic-rich grey sediment filling Zoophycos spreite (white arrows) produced in red pelagic mud (Maastrichtian, Zumaia, Spain); C, homogeneous and pelleted sequestered sediment alternating with host sediment in Zoophycos spreite; inset (C’) showing pellets occurring in both sequestered sediment (green arrows) and host sediment (yellow arrows) (DSDP Leg 93, Site 605, Core 16-4, 59.5-66.5 cm core depth; for details see Wetzel 1987); D, truncation of lamellae (marked with red stippled lines) in Zoophycos spreite in horizontal section, indicating reworking of previously emplaced material (Paleogene, Zumaia, Spain). Scale bars: 1 cm."figureDoi="http://doi.org/10.5281/zenodo.14530602"httpUri="https://zenodo.org/record/14530602/files/figure.png"pageId="5"pageNumber="328">Fig. 1A</figureCitation>
chronometric age data record a sequestration of surface material as the spreite fill is generally younger than the host sediment (e.g.
<bibRefCitationid="EFE34B3FFFEEFF90A903FE5A05F3C50D"author="LOWEMARK L. & WERNER F."pageId="5"pageNumber="328"pagination="191 - 198"refId="ref10979"refString="LOWEMARK L. & WERNER F. 2001. - Dating errors in high-resolution stratigraphy: a detailed X-ray radiograph and AMS- 14 C study of Zoophycos burrows. Marine Geology 177 (3 - 4): 191 - 198. https: // doi. org / 10.1016 / S 0025 - 3227 (01) 00167 - 0"type="journal article"year="2001">Löwemark & Werner 2001</bibRefCitation>
;
<bibRefCitationid="EFE34B3FFFEEFF90AFDBFE7A02E1C50D"author="LEUSCHNER D. C. & SIROCKO F. & GROOTES P. M. & ERLENKEUSER H."box="[967,1195,503,529]"pageId="5"pageNumber="328"pagination="111 - 126"refId="ref10684"refString="LEUSCHNER D. C., SIROCKO F., GROOTES P. M. & ERLENKEUSER H. 2002. - Possible influence of Zoophycos bioturbation on radiocarbon dating and environmental interpretation. Marine Micropalaeontology 46 (1 - 2): 111 - 126. https: // doi. org / 10.1016 / S 0377 - 8398 (02) 00044 - 0"type="journal article"year="2002">
<bibRefCitationid="EFE34B3FFFEEFF90A8ABFE7A03C9C50D"author="KUSSNER K. & SARNTHEIN M. & LAMY F. & TIEDEMANN R."box="[1207,1411,503,529]"pageId="5"pageNumber="328"pagination="48 - 56"refId="ref10553"refString="KUSSNER K., SARNTHEIN M., LAMY F. & TIEDEMANN R. 2018. - High-resolution radiocarbon records trace episodes of Zoophycos burrowing. Marine Geology 403: 48 - 56. https: // doi. org / 10.1016 / j. margeo. 2018.04.013"type="journal article"year="2018">
appears to be produced when environmental conditions switch to a starved sedimentation regime (
<bibRefCitationid="EFE34B3FFFEEFF90AF95FDDA022BC56D"author="KUSSNER K. & SARNTHEIN M. & LAMY F. & TIEDEMANN R."box="[905,1121,598,625]"pageId="5"pageNumber="328"pagination="48 - 56"refId="ref10553"refString="KUSSNER K., SARNTHEIN M., LAMY F. & TIEDEMANN R. 2018. - High-resolution radiocarbon records trace episodes of Zoophycos burrowing. Marine Geology 403: 48 - 56. https: // doi. org / 10.1016 / j. margeo. 2018.04.013"type="journal article"year="2018">
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<bibRefCitationid="EFE34B3FFFEEFF90A8FDFDFB03E2C58C"author="WETZEL A. & TJALLINGII R. & WIESNER M. G."box="[1249,1448,630,656]"pageId="5"pageNumber="328"pagination="256 - 267"refId="ref14786"refString="WETZEL A., TJALLINGII R. & WIESNER M. G. 2011. - Bioturbational structures record environmental changes in the upwelling area off Vietnam (South China Sea) for the last 150,000 years. Palaeogeography, Palaeoclimatology, Palaeoecology 311 (3 - 4): 256 - 267. https: // doi. org / 10.1016 / j. palaeo. 2011.09.003"type="journal article"year="2011">
<bibRefCitationid="EFE34B3FFFEEFF90AF31FD1B0249C5AC"author="DORADOR J. & WETZEL A. & RODRIGUEZ-TOVAR F. J."box="[813,1027,662,688]"pageId="5"pageNumber="328"pagination="323 - 328"refId="ref8866"refString="DORADOR J., WETZEL A. & RODRIGUEZ-TOVAR F. J. 2016. - Zoophycos in deep-sea sediments indicates high and seasonal primary productivity: Ichnology as a proxy in palaeoceanography during glacial-interglacial variations. Terra Nova 28 (5): 323 - 328. https: // doi. org / 10.1111 / ter. 12224"type="journal article"year="2016">
producer stows the sequestered material commonly in the form of mud lamellae or pellets in the spreite. Pellets, however, occur in both host and sequestered sediment, suggesting that they could have provided “seed” microbes housed in the gut to the spreite “bioreactor” (
<figureCitationid="13492A4BFFEEFF90A8DAFCBB0350C44C"box="[1222,1306,822,848]"captionStart="FIG"captionStartId="6.[132,143,1656,1673]"captionTargetBox="[132,1455,215,1609]"captionTargetId="figure-311@6.[603,1266,771,960]"captionTargetPageId="6"captionText="FIG. 1. — Sequestered and downward-transferred material in Zoophycos Massalongo,1855: A, black mud and white nannofossil ooze displaced for> 26 (black arrows) and> 23 cm (white arrows) downward, respectively, as indicated by colour and composition of sediment (Institute of Geological Sciences Kiel, Germany, core 16867-3; 2.2°S, 5.1°E, 3894 m water depth, mid Atlantic; 16.56-16.99 m core depth); B, sequestered organic-rich grey sediment filling Zoophycos spreite (white arrows) produced in red pelagic mud (Maastrichtian, Zumaia, Spain); C, homogeneous and pelleted sequestered sediment alternating with host sediment in Zoophycos spreite; inset (C’) showing pellets occurring in both sequestered sediment (green arrows) and host sediment (yellow arrows) (DSDP Leg 93, Site 605, Core 16-4, 59.5-66.5 cm core depth; for details see Wetzel 1987); D, truncation of lamellae (marked with red stippled lines) in Zoophycos spreite in horizontal section, indicating reworking of previously emplaced material (Paleogene, Zumaia, Spain). Scale bars: 1 cm."figureDoi="http://doi.org/10.5281/zenodo.14530602"httpUri="https://zenodo.org/record/14530602/files/figure.png"pageId="5"pageNumber="328">Fig. 1C</figureCitation>
). Therefore, a priming scenario appears realistic, in particular as subsequent lamellae overlap previous ones, indicating partial reworking and utilization of the spreite fill by the tracemaker (
<figureCitationid="13492A4BFFEEFF90A951FC1803EBC4B3"box="[1357,1441,917,943]"captionStart="FIG"captionStartId="6.[132,143,1656,1673]"captionTargetBox="[132,1455,215,1609]"captionTargetId="figure-311@6.[603,1266,771,960]"captionTargetPageId="6"captionText="FIG. 1. — Sequestered and downward-transferred material in Zoophycos Massalongo,1855: A, black mud and white nannofossil ooze displaced for> 26 (black arrows) and> 23 cm (white arrows) downward, respectively, as indicated by colour and composition of sediment (Institute of Geological Sciences Kiel, Germany, core 16867-3; 2.2°S, 5.1°E, 3894 m water depth, mid Atlantic; 16.56-16.99 m core depth); B, sequestered organic-rich grey sediment filling Zoophycos spreite (white arrows) produced in red pelagic mud (Maastrichtian, Zumaia, Spain); C, homogeneous and pelleted sequestered sediment alternating with host sediment in Zoophycos spreite; inset (C’) showing pellets occurring in both sequestered sediment (green arrows) and host sediment (yellow arrows) (DSDP Leg 93, Site 605, Core 16-4, 59.5-66.5 cm core depth; for details see Wetzel 1987); D, truncation of lamellae (marked with red stippled lines) in Zoophycos spreite in horizontal section, indicating reworking of previously emplaced material (Paleogene, Zumaia, Spain). Scale bars: 1 cm."figureDoi="http://doi.org/10.5281/zenodo.14530602"httpUri="https://zenodo.org/record/14530602/files/figure.png"pageId="5"pageNumber="328">Fig. 1D</figureCitation>
represents very likely a lifetime burrow of its producer (e.g.
<bibRefCitationid="EFE34B3FFFEEFF90AFE9FC5802B0C4F3"author="WETZEL A. & WERNER F."box="[1013,1274,981,1008]"pageId="5"pageNumber="328"pagination="185 - 212"refId="ref14734"refString="WETZEL A. & WERNER F. 1980. - Morphology and ecological significance of Zoophycos in deep-sea sediments off NW Africa. Palaeogeography, Palaeoclimatology, Palaeoecology 32: 185 - 212. https: // doi. org / 10.1016 / 0031 - 0182 (80) 90040 - 1"type="journal article"year="1980">Wetzel & Werner 1980</bibRefCitation>
), the worm-like tracemaker probably took advantage of priming on a time scale of months to years.
