Sequestrichnia - an ethological category of marine trace fossils recording the collection and stowage of nutritional material within burrows
Author
Uchman, Alfred
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)
alfred.uchman@uj.edu.pl
Author
Wetzel, Andreas
Department of Environmental Sciences - Geology, Universität Basel, Bernoullistrasse 32, CH- 4056 Basel (Switzerland) andreas. wetzel @ unibas. ch
andreas.wetzel@unibas.ch
text
Comptes Rendus Palevol
2024
2024-08-30
23
22
325
338
http://dx.doi.org/10.5852/cr-palevol2024v23a22
journal article
305524
10.5852/cr-palevol2024v23a22
fefdf6a6-89e6-4b61-8d87-ff08580d85f9
1777-571X
13595789
urn:lsid:zoobank.org:pub:FAE07554-6F51-46B5-A81D-EA7D9091E776
Ichnogenus
Zoophycos
Massalongo, 1855
(
Fig. 1
)
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 & Gaillard 1996
). In contrast, for Late Cretaceous to modern
Zoophycos
,
sequestrichnial behaviour of the producers appears to be common. These are outlined here.
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 & Uchman 1998
: fig. 1d, e). In the Eocene of Arnakatxa Headland near Bilbao, 0.3-0.7%
Corg
occurs in grey spreite material and <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 & 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
).
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 & Werner 1980
), the worm-like tracemaker probably took advantage of priming on a time scale of months to years.