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FIGURE 1. Location maps: 1, a part of Europe with main localities of the material studied and indication of the main study area in the Polish Carpathians; 2, the main study area in the Polish Carpathians with indication of several localities of the material studied or reported occurrences of Tubulichnium rectum.

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FIGURE 2. Holotype of Tubulichnium rectum (Fischer-Ooster, 1858) and some other key specimens described by Fischer-Ooster (1858) and Książkiewicz (1977), which are housed in the Naturhistorisches Museum der Burgergemeinde Bern (NMBE) in Switzerland and in the Nature Education Centre of the Jagiellonian University - Museum of Geology, Kraków, Poland, respectively: 1, holotype, originally Halymenites rectus Fischer-Ooster and Chondrites targionii (Cht), Gurnigel Flysch (Maastrichtian), Seeligraben near Gurnigelbad (Switzerland), specimen NMBE 5017471. Detail illustrated in 2 marked by the quadrangle; 2, detail of 1; 3, original of Halymenites minor Fischer-Ooster, 1858 and Chondrites intricatus (Chi) from Gurnigel Flysch (Maastrichtian), Seeligraben near Gurnigelbad (Switzerland), specimen NMBE 5017474; 4, original of Halymenites incrassatus Fischer-Ooster, 1858, Fähnernspitz, Upper Cretaceous, E Switzerland, specimen NMBE 5017470; 5, holotype of Tubulichnium incertum Książkiewicz, 1977 (yellow arrow), other specimen of the same ichnotaxon (Tr) and Scolicia vertebralis (Sv); Ropianka Formation (Inoceramian Beds), Upper Cretaceous-Paleocene, Bachów, Skole Nappe, specimen UJTF 938; 6, detail of 5 showing the holotype.

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FIGURE 3. Tubulichnium rectum (Fischer-Ooster, 1858) on upper bedding surfaces from different formations: 1, on the upper surface of turbiditic marl with Chondrites intricatus (Chi), Pagliaro Formation (Paleocene), Cabella Ligure, Northern Apennines, Italy, field photograph; 2, curved example of T. rectum (Tr), Scolicia isp. (Sc) and Phycosiphon incertum (Ph), Ropianka Formation (Upper Cretaceous), Wara, Skole Nappe, Carpathians, Poland, UJTF 1426, Książkiewicz collection; 3, several specimens of T. rectum (Tr), some cross cut by Chondrites intricatus (Chi), Pagliaro Formation (Paleocene), Cabella Ligure, Northern Apennines, Italy, field photograph; 4, specimen with two (1, 2) superimposed pellet pavements. Inoceramian Beds (Upper Cretaceous-Paleocene), Słopnice, INGUJ144P190a; 5; specimens with elevated edges and collapsed roof, Inoceramian Beds (Upper Cretaceous-Paleocene), Słopnice, INGUJ144P140; 6, example of spindle-shaped burrow enlargement, Pagliaro Formation (Paleocene), Cabella Ligure, Northern Apennines, Italy, field photograph; 7, T. rectum cut by Ophiomorpha annulata (Oa), Inoceramian Beds (Upper Cretaceous-Paleocene), Słopnice, INGUJ143P66.

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FIGURE 4. Tubulichnium rectum (Fischer-Ooster, 1858) on the lower bedding surfaces (1-3) of and in longitudinal (4) and transverse (5-10) cross-sections of turbiditic sandstones from the Inoceramian Beds (Upper-Cretaceous-Paleocene at Słopnice: 1, note external and internal parts INGUJ144P176; 2, note sandy mantle covering the pelleted part, INGUJ144P193a, 3, note wrinkles on the mantle, INGUJ144P187a; 4, longitudinal cross section (Tr), INGUJ144P193; 5-10, transverse sections, E - INGUJ144P182d, F - INGUJ144P180, G - INGUJ144P192b, H - INGUJ144P170a, I - INGUJ144P190, J - INGUJ144P192c, K - INGUJ144P192.

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FIGURE 5. Macroscopic and microscopic and SEM observations of Tubulichnium rectum (Fischer-Ooster, 1858) occurring in turbiditic sandstones of the Inoceramian Beds (Upper Cretaceous-Paleocene), Słopnice, Poland (1, 4-8) and turbiditic marls of the Pagliaro Formation (Paleocene), Italy (2, 3): 1, specimen with elevated edges and bent down roof as seen on upper bedding surface (INGUJ144P195); 2, 3, depressions of pellets in the burrow margin after pellets have been partly or completely removed (INGUJ196P36 and 38); 4, photomicrograph of transverse cross-section; note deformed laminae above the collapsed burrow; 5, 6, burrow margin showing depressions of pellets that have been removed by weathering; 7, part of split pellets in place consisting of clay minerals and siliciclastic silt grains; 8, outer part of a split pellet showing platy minerals oriented ±parallel to surface.

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FIGURE 6. Short sections showing position of Tubulichnium rectum and associated traces fossils in strata of the Ropianka Formation (Inoceramian Beds) at Słopnice, Magura Nappe, Carpathians, Poland. GPS coordinates: section A: 49°42.982'N, 020°20.448'E; section B: 49°42.974'N, 20°20.453'E, section C: 49°42.610'N, 20°20.687'E.

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FIGURE 7. Stratigraphic range of Tubulichnium rectum (Fischer-Ooster, 1858) and number of formations per stage in which it occurs.

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TABLE 1. Occurrences of Tubulichnium rectum (Fischer-Ooster, 1858). * - reserve inclusion in T. rectum; ** - probably not T. rectum.

Formation and general location Localities Age Facies References
Al Ayn Formation, Oman   Upper Triassic Deep-sea turbiditic sandstones and shales Wetzel et al. (2007)
Globotruncana Marl, Pieniny Klippen Belt, Carpathians, Poland Sromowce Wyżne Turonian Deep-sea, pelagic and hemipelagic sediments Książkiewicz (1977)
Zementmergelserie, Rhenodanubian Flysch, Bavarian Alps, Germany Kalkgraben quarry, Arzbach Stream, Steinbach near Benediktbeuern, Halbammer Valley Turonian-Maastrichtian Turbiditic limestones, marlstones and mudstones Uchman (1999)
Sromowce Beds, Pieniny Klippen Belt, Carpathians, Poland Sromowce Wyżne Coniacian-Maastrichtian Deep-sea turbiditic sandstones and shales Książkiewicz (1977)
Holovnia Marl, Skole Nappe Carpathians, Poland Krzeczkowa - old quarry Coniacian-Santonian Deep-sea marls This study
Ropianka Formation (Inoceramian Beds), Skole Nappe, Carpathians, Poland, Ukraine Poland: Wara, Wołodź, Kuźmina, Bachów, Huwniki, Kalwaria Pacławska (the steram Sopotnik), Wola Romanowa. Ukraine: Dora (Yaremche) Coniacian-Paleocene Deep-sea turbiditic sandstones and marls Książkiewicz (1977); this study
Kropivnik Fucoid Marls, Skole Nappe, Carpathians, Poland Wiar section Campanian- lower Maastrichtian Deep-sea marls Leszczyński (2004)
Tuscan Apennines, Italy Alberese Campanian-Eocene Deep-sea marls Reis (1909); Abel (1935); Seilacher (1959)
Monte Antola Formation, Ligurian Domain, Northern Apennines, Italy Vagge, Molino Vecchio, Val Gardenella, Monte Colletto, Candini, Roiale Late Campanian - Maastrichtian Deep-sea turbiditic marls, sandstones and shales Uchman (2007b, 2009)
Northpenninic Flysch, Alps, E Switzerland Fähnernspitz Mt. Upper Cretaceous Deep-sea turbiditic marls, sandstones and shales Fischer-Ooster (1858)
Ropianka Formation, Magura Nappe, Carpathians, Poland Złatna, Mordarka, Biczyce, Wola Brzezińska, Wola Krogulecka, Siary, Uście Gorlickie, Koninki, Stara Wieś, Lubomierz, Słopnice Maastrichtian-Paleocene Deep-sea turbiditic sandstones, shales, locally marls Książkiewicz (1977); Uchman (1991c, 1998); this study
“Zumaya Flysch”, Basque Country, Spain No data Upper Cretaceous - lower Eocene Deep-sea turbiditic limestones, marls, sandstones and shales Cummings and Hodgson (2011)*
Upper Istebna Beds, Silesian Nappe, Carpathians, Poland Kamesznica (the stream Janoska) Paleocene Deep-sea turbiditic sandstones and shales Książkiewicz (1977)
Kanina Beds, Magura Nappe, Carpathians, Poland Lubomierz Paleocene Deep-sea turbiditic marls and shales Cieszkowski et al. (1992)
Itzurun Formation, Basque Country, Spain Zumaya section Paleocene Deep-sea turbiditic Limestones, marls, sandstones and shales Giannetti (2010)
Pagliaro Formation, Ligurian Domain, Northern Apennines, Italy Pallavicino, Celio, Salata, Cabella Ligure, Fubbiano, Borassi Paleocene Deep-sea turbiditic sandstones, marls and shales Uchman (2007a); this study
Greifenstein Sandstone, Wienerwald, Rhenodanubian Flysch, Austria Hof bei Karsten, Schleifsteinbruch Eocene Deep-sea turbiditic sandstones and shales Abel (1935); Uchman (1999); Kern (1978)
Variegated Shale, Magura Nappe, Carpathians, Poland Wola Brzezińska Upper Paleocene-Upper Eocene Deep-sea shales with turbiditic sandstones Książkiewicz (1977)
Szczawnica Formation, Magura Nappe, Carpathians, Poland Hałuszowa, Krościenko-Łąkcica, Szczawnica, Szlachtowa, Łomnica, Wierchomla Mała, Łosie, Jamne, Jaszcze, Rzyczanów Upper Paleocene-Lower Eocene Deep-sea turbiditic sandstones and shales Książkiewicz (1977); Uchman (1991c, 1992a, 2008a); Wetzel and Uchman (1998); this study
Gurnigel Flysch, Fribourgian Alps, Switzerland Seeligraben n. Gurnigelbad, Zollhaus Upper Paleocene Deep-sea, pelagic and hemipelagic sediments Fischer-Ooster (1858); Crimes et al. (1981), Uchman (1998, figure 26B, C)
Flysch del Grivó, Julian Prealps, NE Italy Clapp, Vernasso Upper Paleocene-Lower Eocene Deep-sea turbiditic sandstones and shales Tunis and Uchman (1993, 1996)
Beloveža Formation, Magura Nappe, Carpathians, Poland Sidzina, Zbludza Lower-Middle Eocene Deep-sea turbiditic sandstones and shales Książkiewicz (1977); this study
Hieroglyphic Beds, Magura Nappe, Carpathians, Poland Rajcza (?) Eocene Deep-sea turbiditic sandstones and shales Książkiewicz (1977)
Hieroglyphic Beds, Dukla Nappe, Carpathians, Poland Daliowa Eocene Deep-sea turbiditic sandstones and shales Książkiewicz (1977)
Piwniczna Sandstone Member of the Magura Formation, Carpathians, Poland Tylmanowa-Baszta Lower Eocene Deep-sea Turbiditic sandstones and shales This study
Bisericani Formation, Marginal Folds Unit, Eastern Carpathians, Romania Mt. Geamana, Valea Corbu Upper Eocene Marls Krejci-Graf (1936)
Numidian Flysch, Atlas, Tunisia Meloula-Tabarka Upper Oligocene Thin-bedded turbidites interbedded with light green mudstones Riahi et al. (2014)*
Middle Bhuban molasse sediments of Mizoram, Northeast India No data Miocene Intertidal to subtidal clastics Rajkumar et al. (2012)**
 

uchmanAlfred Uchman. Institute of Geological Sciences, Jagiellonian University, Gronostajowa 3a, PL 30-387 Kraków, Poland; This email address is being protected from spambots. You need JavaScript enabled to view it.

Alfred Uchman is an ichnologist working on invertebrate ichnology in marine and non-marine facies, mostly in Europe. His interest is focused mostly on ichnotaxonomy and palaeoecology of trace fossils in deep-sea sediments. He graduated from the Jagiellonian University in Kraków, Poland. Since that time he is working in the Institute of Geological Sciences of the same university. Alfred Uchman is the titular professor since 2001, a member of the Polish Academy of Art and Sciences and the Polish Academy of Sciences. In 2012-2016 he was the President of the International Ichnological Society.

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wetzelAndreas Wetzel. Geologisches Institut, Universität Basel, Bernoullistrasse 32, CH-4056 Basel, Switzerland; This email address is being protected from spambots. You need JavaScript enabled to view it.

Andreas Wetzel is sedimentologist working on Holocene and Pleistocene sediments in the South China Sea and also in Mesozoic sedimentary rocks in Switzerland and Argentina. Since his PhD at the University in Kiel (Germany) on bioturbation processes in Late Quaternary deep-sea sediments off NW Africa he is continuously interested in ichnology. From 1980 to 1987 he worked as postdoc and assistant professor at the University of Tübingen (Germany) on physical properties of deep-sea sediments focusing on DSDP and ODP material. Since 1987 he is professor at the University of Basel (Switzerland) teaching sedimentology. 

 
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Hidden subsurface garden on own faeces – the trace fossil Tubulichnium rectum (Fischer-Ooster, 1858) from the Cretaceous-Palaeogene deep-sea sediments

Alfred Uchman and Andreas Wetzel

Plain Language Abstract

In the deep-sea, food for benthic organisms is mainly provided by organic matter produced in the photic zone that settles to the seafloor. Due to large water depth, the settling time of organic matter is long and hence, a high proportion of this material becomes oxidized. Thus, the deep-sea floor represents a food-limited habitat. Consequently, a specialized nutritional strategy is essential for long-term survival. To ingest organic-rich material that arrived after a bloom in surface water on the seafloor and to store it in anoxic sediment for later utilization is a successful strategy as described in this study.

Resumen en Español

Reserva oculta en el subsuelo con las propias heces ¬– la traza fósil Tubulichnium rectum (Fischer-Ooster, 1858) de los sedimentos marinos profundos del Cretácico-Paleógeno

Tubulichnium rectum (Fischer-Ooster, 1858) es un tubo, entre oblicuo y horizontal, no ramificado, con terminación cerrada, que tiene márgenes densamente alineados con pellets elipsoidales de naturaleza fangosa. Se presenta en sedimentos marinos profundos de arena fina a fango, siliciclásticos y margosos, principalmente de Turoniense al Eoceno. Probablemente fue producido por organismos semejantes a "gusanos", los cuales se alimentaban de sedimentos ricos en materia orgánica depositados de manera estacional o episódica en el fondo marino. Los pellets fecales se concentraban en las partes profundas de la traza dentro de la zona anóxica más allá de la profundidad a la que usualmente penetran otros excavadores. Durante períodos de escasez de alimento, los pellets se utilizaron como una fuente de nutrición suplementaria. Este comportamiento de acumulación y uso de unas reservas se interpreta como una adaptación a la creciente competencia por los alimentos en el mar profundo después del Cenomaniense.

Palabras clave: icnología; icnotaxonomía; autocoprofagia; cache; flysch; turbiditas

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Tubulichnium rectum (Fischer-Ooster, 1858) est un tube non branché, oblique à horizontal en cul-de-sac. Il possède une périphérie alignée avec pellets boueux ellipsoidales. Cet ichnotaxon est repartie du Turonian jusq'à l'Eocène et se retrouve dans des sediments marins sableux, siliclastiques et marneux de grand profondeur. Il est produit par des organismes vermiculaires qui se nourrissait de sediment riche en matière organique déposé saisonièrement ou occasionellement sur le fond marin. Les pellets fécales étaient gardés dans le tube plus profond que les terriers' qui pénètre la couche anoxique du sédiment. Pendant des périodes de pénurie alimentaires les organismes utilisaient les pellets comme source de nourriture. Le comportment de construire des dépots alimentaires est interprété comme adapdation à une competition augmentée dans l'abyssale après le Cénomanien.

Translator: Christian Meyer

Deutsche Zusammenfassung

Versteckte unterirdische Gärten auf eigenen Kot-Ausscheidungen – das Spurenfossil Tubulichnium rectum (Fischer-Ooster, 1858) in kretazischen bis paläogenen Tiefsee-Sedimenten

Tubulichnium rectum (Fischer-Ooster, 1858) ist ein schräg bis horizontal im Sediment verlaufender, unverzweigter, blind endender Gang, an dessen Rand sich dicht gepackt elliposidförmige pelitische Kotpillen befinden. Diese Spur tritt in feinsandigen bis pelitischen siliziklastischen und mergeligen Tiefsee-Sedimenten auf, vor allem vom Turon bis zum Eozän. Dieses Spurenfossil wurde wohl von einem wurmförmigen Organismus erzeugt, der organisch-reiches Material aufnahm, das saisonal oder episodisch auf der Meeresbodenoberfläche abgelagert wurde. Es wurde in Form von Kotpillen im Gang in anoxischem Sediment in so grosser Tiefe ausgeschieden, dass es sich ausserhalb der Reichweite anderer Organismen befand. Während Zeiten geringer Nahrungsverfügbarkeit wurden die Kotpillen vermutlich als zusätzlich Nahrung genutzt. Dieses Verhalten, in einem Lager nahrhaftes Material zu speichern, wird als Anpassung der erzeugenden Organismen an ein Habitat gesehen, in dem seit dem Cenoman eine zunehmende Konkurrenz zwischen benthischen Organismen um Nahrung besteht.

Translator: Andreas Wetzel

Arabic

Translator: Ashraf M.T. Elewa