identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03DB87D8FFEEFF90AF81FBD8034FC010.text	03DB87D8FFEEFF90AF81FBD8034FC010.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Polykampton Ooster 1869	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Polykampton Ooster, 1869</p>
            <p> Polykampton occurring in Cretaceous (Albian) to Oligocene deposits represents a horizontal median tunnel with lateral spreite lobes, which alternate on either side of the tunnel and are slightly inclined to bedding; the tunnel runs within the interval of the spreite lobes or underneath (Uchman et al. 2019).  Polykampton is produced: 1) in the sandy or muddy interval of (turbiditic) event beds; 2) along the mudstonesandstone interface; or 3) across the sandstone-mudstone transition in turbiditic beds. The spreite and the main tunnel contain mud that is enriched in organic matter compared to the surrounding host sediment (Corg 3.05% vs 1.05%; Uchman et al. 2019). The mud was evidently introduced from above into the burrow. The organic content of spreite lobes could be a source of additional benthic food in the short term, just after organic-rich material was sequestered on the surface and stowed in spreite lobes. In addition, or alternatively, the producer could utilize organic matter when passing through the tunnel during the construction of a new lobe. By then, the sequestered organic-rich material might have already been altered to some degree by microbial activity. </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFEEFF90AF81FBD8034FC010	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFEEFF90ACF5FB19027DC333.text	03DB87D8FFEEFF90ACF5FB19027DC333.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zoophycos Massalongo 1855	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Zoophycos Massalongo, 1855</p>
            <p>(Fig. 1)</p>
            <p> Zoophycos 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. Seilacher 1977b; later refined by Chamberlain 2000 and Zhang et al. 2015). For most  Zoophycos older than the late Mesozoic, different ways of spreite production and behaviour are possible, including fodinichnial behaviour (e.g. Olivero &amp; Gaillard 1996). In contrast, for Late Cretaceous to modern  Zoophycos , sequestrichnial behaviour of the producers appears to be common. These are outlined here. </p>
            <p> 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 types of lamellae (Fig. 1). 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  Zoophycos (Gurnigel Flysch; Seligraben/Gurnigelbad, Switzerland) contains 1.1-1.7% Corg compared to 0.5-0.7% Corg in the green host sediment (see Wetzel &amp; Uchman 1998: fig. 1d, e). In the Eocene of Arnakatxa Headland near Bilbao, 0.3-0.7% Corg occurs in grey spreite material and &lt;0.1% Corg in red limestone lutite alternations. However, in deep oceanic settings, also lighter material deposited during periods of enhanced nannoplankton productivity can be transferred downward (Fig. 1A). Other tracers, e.g. volcanic ash, also record a downward sediment transfer (e.g. Kotake 1991). For Pleistocene and Holocene  Zoophycos , chronometric age data record a sequestration of surface material as the spreite fill is generally younger than the host sediment (e.g. Löwemark &amp; Werner 2001; Leuschner et al. 2002; Küssner et al. 2018). In well-dated sediments,  Zoophycos appears to be produced when environmental conditions switch to a starved sedimentation regime (Küssner et al. 2018) or during times of enhanced seasonality (e.g. Löwemark et al. 2006; Wetzel et al. 2011; Dorador et al. 2016). </p>
            <p> The  Zoophycos 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” (Fig. 1C). 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 (Fig. 1D). Since  Zoophycos represents very likely a lifetime burrow of its producer (e.g. Wetzel &amp; Werner 1980), the worm-like tracemaker probably took advantage of priming on a time scale of months to years. </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFEEFF90ACF5FB19027DC333	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFEEFF92AF6BF8BF07A1C492.text	03DB87D8FFEEFF92AF6BF8BF07A1C492.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tubulichnium Ksiazkiewicz 1977	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Tubulichnium Książkiewicz, 1977 (Fig. 2) </p>
            <p> Tubulichnium is an oblique to horizontal, unbranched, blindending tube showing some internal organization due to slight vertical shift; the margins are densely lined with ellipsoidal muddy pellets. The sequestration of material, which is then stowed in muddy laminae and pelleted fill, is documented by its composition: it contains c. 1.5% Corg and 20% CaCO 3 compared to c. 1.1% Corg and c. 26% CaCO 3 in the host sediment and 0.7% Corg and 12% CaCO 3 in the overlying mud in the studied Upper Cretaceous to Paleogene specimens (Uchman &amp; Wetzel 2017). These data point to selective enrichment of organic-rich particles in the burrow. Microbial activity during priming may generate CO 2 that lowers alkalinity and fosters the dissolution of carbonate (van Nugteren et al. 2009). Although no clear spreite is developed, a partly laminated fill exhibiting local truncations (Fig. 2B, C) suggests utilization of the stowed material, as in the case of  Zoophycos . </p>
            <p>TUBULAR OPEN BURROW SYSTEMS</p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFEEFF92AF6BF8BF07A1C492	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFECFF92AC86FC380735C030.text	03DB87D8FFECFF92AC86FC380735C030.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cladichnus D'Alessandro & Bromley 1987	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Cladichnus D’Alessandro &amp; Bromley, 1987</p>
            <p> The  Cladichnus burrow (Cretaceous-Eocene) consists of primary successively branched and radiating tubes, which contain a meniscate fill. These traces are preferentially constructed in anoxic sediments. New branches are produced roughly at the same level as previous ones, implying that appropriate conditions for stowing were encountered by the producer. In contrast, new branches constructed below or above may target stronger or weaker reducing conditions or represent a response to the downward or upward migration of the redox boundary. </p>
            <p> Collection and transfer of surface material is recorded by the tubes’ fill that was analyzed in detail for the ichnospecies  Cladichnus parallelum Wetzel &amp; Uchman, 2013 , which contains 0.8% Corg and 0.3% CaCO 3 compared to 0.3% Corg and 64% CaCO 3 in the host sediment Cretaceous in age (Wetzel &amp; Uchman 2013). The fill of the branches is interpreted to indicate priming that provided microbes (or their metabolic products) as an additional subsurface food source (cf. Mayer et al. 2001; van Nugteren et al. 2009). </p>
            <p>Most likely, the trace producer ingested a considerable proportion of the filling material before emplacing the final, now meniscate fill. The arrangement of the branches has further implications. Branches at a level above the previous ones could allow utilization of upstreaming pore water, probably carrying nutritious compounds. Alternatively, a shift of the redox boundary may have caused the construction of new levels of open burrows.</p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFECFF92AC86FC380735C030	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFECFF92AF98FBD8027CC1D1.text	03DB87D8FFECFF92AF98FBD8027CC1D1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ichnogenus Thalassinoides Ehrenberg 1944	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus Thalassinoides Ehrenberg, 1944</p>
            <p>(Fig. 3)</p>
            <p>Crustaceans living in the modern deep South China Sea produce Thalassinoides- like burrows that document a sequestrichnial behaviour. Two kinds of burrows were encountered, which contained abundant foraminiferal tests stowed in greenish anoxic sediment. One burrow type contains benthic agglutinated foraminifera tests, which consist mainly of Pinatubo-1991 ash but were stowed below the ash in anoxic sediment (Kaminski &amp; Wetzel 2004). The other kind of Thalassinoides occurs in water depths below the CCD; it exhibits a laminated fill consisting of calcareous planktonic foraminiferal tests that are stowed and preserved in the burrow, whereas they are already dissolved in the overlying hemipelagic sediment from which they originate (Wetzel &amp; Unverricht 2013). The calcareous foraminiferal tests must have been stowed shortly after their deposition; otherwise they would show dissolution features. Partly truncated laminae in the burrow imply later reworking by the producer (Fig. 3).</p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFECFF92AF98FBD8027CC1D1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFECFF92ACC8F8DF0280C333.text	03DB87D8FFECFF92ACC8F8DF0280C333.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phymatoderma Brongniart 1849	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Phymatoderma Brongniart, 1849</p>
            <p> Phymatoderma (Lower Jurassic-Pliocene) appears to be similar in some aspects to  Cladichnus since it represents an actively filled burrow system consisting of numerous branches, which deviate at a few levels from a common stem, diverging distally (e.g. Fu 1991; Izumi 2012). However, the producers tended to operate in a plane or only a few levels, less than in  Cladichnus . In most cases, the branches are filled with pellets that are darker than the surrounding material (e.g. Uchman 1999), suggesting a higher content of organic matter that could foster enhanced microbial activity (e.g. Izumi et al. 2015). In black shales, in contrast, the pellets are lighter than the host sediment but document incorporation of surface detritus (Seilacher 1978; Izumi 2012). </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFECFF92ACC8F8DF0280C333	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
03DB87D8FFECFF9CAF47F97F07DBC3D2.text	03DB87D8FFECFF9CAF47F97F07DBC3D2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Avetoichnus Uchman & Rattazzi 2011	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> Ichnogenus  Avetoichnus Uchman &amp; Rattazzi, 2011 (Fig. 4) </p>
            <p> Avetoichnus (Paleocene-Oligocene) represents a mid-tier burrow consisting of a mostly horizontal to subhorizontal helix enveloping a central tube that was probably an open tunnel when the tracemaker lived. The tube contains grey mud, whereas the spiral is filled with black, probably organic-rich mud that likely originated from the overlying pelagic sediment (Uchman &amp; Rattazzi 2011). The tube resembles a pinched open burrow that truncates the inner edges of the helical turns, which were already filled with dark mud when the tunnel was still inhabited. Chemichnia like  Chondrites co-occur with  Avetoichnus (Fig. 4). </p>
            <p>The colour of the helical fill suggests that it was displaced downward. Furthermore, the partial truncation of the inner turns of the spiral points to later reworking by the tracemaker. Very likely, the organic material in the spiral fostered the development of reducing conditions therein whereas the central tunnel was open and oxygenated water could be circulated through it by the inhabitant. Thus, a steep geochemical gradient developed across the tunnel margin between anoxic host sediment and oxic water in the lumen, a situation known to enhance microbial activity (e.g. Meysman et al. 2010). Thus, the trace-fossil producer sequestering organic-rich material on the seafloor appears to have constructed an organic-rich subsurface domain that rapidly became anoxic. Therefore, this trace is classified as a sequestrichnion.</p>
            <p>OTHER POSSIBLE SEQUESTRICHNIA</p>
            <p>The burial of labile organic matter in anoxic sediments deep below the surface mixed layer strongly suggests priming induced by the trace producers, probably wormlike animals such as echiurans, sipunculids, and polychaetes together with crustaceans. Besides utilizing the sequestered burrow fill directly as indicated by subsequent reworking, some trace-fossil producers burrowed down and construct a cache underneath a domain, which is already enriched in organic matter like seagrass or wood (e.g. Griffis &amp; Suchanek 1991; Bojanowski &amp; Wetzel 2024). From this perspective, some other trace fossils could also be sequestrichnia.</p>
            <p> At least some specimens of the spreite burrow Teichichnus  zigzag Frey &amp; Bromley, 1985 occurring in Jurassic deposits record sequestrichnial behaviour (e.g. Wetzel et al. 2023). Teichichnus  zigzag exhibits: 1) a spreite enriched in organic-rich material relative to the host sediment; 2) no reworking halo around the spreite (thus indicating that the organic matter was introduced from above); 3) lamellae produced later that systematically crosscut previously produced ones, and hence indicate reworking by the producer. It can thus be excluded that reworking represents a response to erosion of the sediment surface, because reworking of the spreite is locally restricted. Furthermore, in calm lagoonal bayfill deposits,  T. zigzag is common (Knaust 2018). In these settings, hyperpycnal flows may provide organic matter. Therefore, a sequestrichnial behavior of the  T. zigzag tracemakers in calm lagoonal settings is highly likely though it has not yet been demonstrated. </p>
            <p> Other spreite burrows, for instance  Rhizocorallium commune Schmid, 1876 , are extensively filled with pellets. They were produced in dysoxic, low-energy mud (Knaust 2013), and could represent sequestrichnia. Unfortunately, TOC data of the spreite fill and host sediment are not available and, therefore, clear evidence of sequestrichnial behaviour is lacking although it is not unlikely. </p>
            <p> Halimedides Lorenz von Liburnau, 1902 , Cambrian (Series 2) to Recent in age, is a system of horizontal tunnels with chambers strung along it, not rarely in the context of poorly oxygenated deposits (Fernández-Martínez et al. 2021). When preserved in full relief, the chambers are filled with darker material than in the surrounding rock (e.g. Uchman 1999). The chambers, which have been interpreted as food caches (Gaillard &amp; Olivero 2009; Lukeneder et al. 2012), could be good places for microbial processing of organic matter.  Lepidenteron mantelli (Geinitz, 1850) is a burrow in Upper Cretaceous marls of the deeper continental shelf. It is filled with sediment rich in plant material, which is strongly pyritized, foremost with pyrite framboids (Jurkowska et al. 2018). The plant material was actively collected, was a substrate for microbial activity and probably promoted priming. Therefore, this trace fossil could be partly sequestrichnial and partly chemichnial.  Gyrophyllites Glocker, 1841 , Cambrian(?) and Early Ordovician to Cenozoic in age, is composed of a vertical to subvertical shaft with lateral, radial, petal-like lobes that may be repeatedly distributed at a few levels. The lobes may be filled with organic-rich material from the underlying or overlying bed, and their position at different levels was modulated by the migration of the redox boundary (Strzeboński &amp; Uchman 2015; Muñoz et al. 2019). The fill of the lateral lobes could promote microbial processing including priming. However, in the case of  Gyrophyllites , the mud appears to have been taken not only from the seafloor but also from of a bed beneath it, in cases where the proper mud was not present on the seafloor, in particular if a decelerating, low-erosive gravity flow rapidly deposited sand that covered and preserved the previous seafloor including the topmost organic-rich interval. Some  Planolites Nicholson, 1873 from Cretaceous marls was evidently filled actively with darker sediment from above (Locklair &amp; Savrda 1998), possibly to induce priming. As  Planolites can be produced by diverse organisms in a wide range of environments, only certain representatives of this ichnogenus might be sequestrichnia. Also,  Alcyonidiopsis Massalongo, 1856 , a simple, cylindrical burrow filled with pellets, commonly darker than the surrounding deposits, and known at least since the Ordovician (see Uchman 1999) can belong to sequestrichnia. </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/03DB87D8FFECFF9CAF47F97F07DBC3D2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Uchman, Alfred;Wetzel, Andreas	Uchman, Alfred, Wetzel, Andreas (2024): Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows. Comptes Rendus Palevol 23 (22): 325-338, DOI: 10.5852/cr-palevol2024v23a22, URL: http://dx.doi.org/10.5852/cr-palevol2024v23a22
