Phosphatized bromalites from the lower Paleozoic of NW Argentina

 

Authors: Voldman GG, Zeballo FJ, Vaccari NE, Carrera MG, Knaust D, Hunt AP

Published in: Bulletin of Geosciences, volume 101, issue 1; pages: 3 - 18; Received 4 February 2025; Accepted in revised form 15 September 2025; Online 9 November 2025

Keywords: feces, trace fossil, paleoecology, Cambrian, Ordovician,

full text (PDF, 14.46 MB)

Export to RIS

 

Abstract

Bromalites, the fossilized remains of material sourced from the digestive system of organisms, are often overlooked in micropaleontological samples, but offer valuable insights into the paleoecology of ancient ecosystems. This study focuses on Furongian (upper Cambrian) to Floian (Lower Ordovician) bromalites from the Central Andean Basin of the Eastern Cordillera, NW Argentina. In many samples, bromalites dominate as the primary organic residue after acid treatment. The recovered assemblage consists of 276 coprolites and cololites, including Anfractuosocoprolithus aphelus Bischoff, A. cf. aphelus Bischoff, Coprulus oblongus Mayer, Hyperocoprolithus levigatus Bischoff, Isvilina kanygini Schallreuter, Rhizocorallites articularis Müller, Transmeristocoprus multifidus Bischoff, Transmeristocoprus? isp., and three unidentified ichnospecies (A, B, and C). Furthermore, a new ichnogenus, Helicobromites, is established, with the description of a new ichnospecies, H. andinus. These ichnospecies may reflect the activity of mollusks as well as that of poorly documented organisms such as ascidian tunicates, polychaetes, holothurians, and perhaps nektonic vertebrates. In this way, the coprolites and cololites from the Eastern Cordillera significantly enhanced the reconstruction of lower Paleozoic ecosystems in the southwestern Gondwanan seas

References

Acenolaza, G.F. 1996. Bioestratigrafía del límite Cámbrico-Ordovícico y Ordovícico basal en la Quebrada de Humahuaca, Provincia de Jujuy, República Argentina. 245 pp. Master thesis, Facultad de Ciencias Naturales e IML, Universidad Nacional de Tucumán, Argentina.

Acenolaza, G.F., Sá, A.A. & Gutiérrez-Marco, J.C. 2008. Cruziana yini Yang, a peri-Gondwanan trilobite trace with new records in the Ordovician of South America and Iberia. Cuadernos del Museo Geominero 9, 19-26.

Aldridge, R.J., Gabbott, S.E., Siveter, L.J. & Theron, J.N. 2006. Bromalites from the Soom Shale Lagerstätte (Upper Ordovician) of South Africa: palaeoecological and palaeobiological implications. Palaeontology 49(4), 857-871. View article

Arakawa, K.Y. 1971. Studies on the faecal pellets of marine invertebrates (excluding molluscs) I. Publications of the Seto Marine Biological Laboratory 19, 231-241. View article

Aráoz, L. 2009. Microfloras ordovícicas en Sierra de Zenta, Cordillera Oriental Argentina. Serie Correlación Geológica 25, 37-94.

Argyriou, T., Clauss, M., Maxwell, E.E., Furrer, H. & Sánchez Villagra, M.R. 2016. Exceptional preservation reveals gastrointestinal anatomy and evolution in early actinopterygian fishes. Scientific Reports 6, 18758. View article

Astini, R.A. 2003. The Ordovician Proto-Andean basins, 1-74. In Benedetto, J.L. (ed.) Ordovician fossils of Argentina. Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba.

Astini, R.A. 2008. Sedimentación, facies, discordancias y evolución paleoambiental durante el Cámbrico-Ordovícico, 50-73. In Coira, B. & Zappettini, E.O. (eds) Geología y Recursos Naturales de Jujuy Relatorio del XVII Congreso Geológico Argentino. Asociación Geológica Argentina, Buenos Aires.

Astini, R.A., Waisfeld, B.G., Toro, B.A. & Benedetto, J.L. 2004. El Paleozoico inferior y medio de la región de Los Colorados, borde occidental de la Cordillera Oriental (provincia de Jujuy). Revista de la Asociación Geológica Argentina 59(2), 243-260.

Bandel, K. 1974. Faecal pellets of Amphineura and Prosobrandria (Mollusca) from the Caribbean coast of Columbia, South America. Senckenbergiana Maritima 6(1), 1-31.

Bertling, M., Buatois, L.A., Knaust, D., Laing, B., Mángano, M.G., Meyer, N., Mikuláš, R., Minter, N.J., Neumann, C., Rindsberg, A.K., Uchman, A. & Wisshak, M. 2022. Names for trace fossils 2.0: theory and practice in ichnotaxonomy. Lethaia 55, 1-19. View article

Bicknell, R.D.C., Smith, P.M. & Kimmig, J. 2023. Novel coprolitic records from the Silurian (Přídolí) Wallace Shale of New South Wales. Alcheringa 47(1), 24-30. View article

Bicknell, R.D.C., Bruthansová, J. & Kimmig, J. 2024. Shelly coprolites record durophagous predation in the Late Ordovician Bohdalec Formation (Katian; Prague Basin, Czech Republic). Geobios 82, 1-11. View article

Bischoff, G.C.O. 1990. Coprolites from Early and Middle Silurian strata of New South Wales. Senckenbergiana lethaea 70(1-3), 209-237.

Bischoff, G.C.O. 1994. Early Ordovician (Arenigian) coprolites from Australia, partly composed of Arenigiapalla emanuela n. g., n. sp., a phosphatised nannofossil. Courier Forschungsinstitut Senckenberg 169, 275-297.

Buatois, L.A., Zeballo, F.J., Albanesi, G.L., Ortega, G., Vaccari, N.E. & Mángano, M.G. 2006. Depositional environments and stratigraphy of the Upper Cambrian-Lower Ordovician Santa Rosita Formation at the Alfarcito area, Cordillera oriental, Argentina: integration of biostratigraphic data within a sequence stratigraphic framework. Latin American Journal of Sedimentology and Basin Analysis 13(1), 1-29.

Buchholz, A. 2015. An uncommon bed structure in a glacial erratic of Cambrian from Western Pomerania (Northeastern Germany) - lithology and fauna. Geschiebekunde aktuell 31(1), 11-22.

Carlorosi, J. & Heredia, S. 2013. The conodont Trapezognathus diprion (Lindström) in the Acoite Formation, Eastern Cordillera, northwestern Argentina, 1-4. In Albanesi, G.L. & Ortega, G. (eds) Conodonts from the Andes, Proceedings of the 3rd International Conodont Symposium, Asociación Paleontológica Argentina, Publicación Especial 13. Buenos Aires.

Castellani, C., Maas, A., Eriksson, M.E., Haug, J. T., Carolin Haug, C. & Waloszek, D. 2018. First record of Cyanobacteria in Cambrian Orsten deposits of Sweden. Palaeontology 61(6), 855-880. View article

Cosma, T.N. & Baumiller, T.K. 2005. A Trace Fossil on a Silurian Bivalve: Evidence of Predatory Boring? Ichnos 12(2), 135-139. View article

Dattilo, B.F., Freeman, R.L., Peters, W.P., Heimbrock, W.P., Deline, B., Martin, A.M., Kallmeyer, J.W., Reeder, J. & Argast, A. 2016. Giants among micromorphs: were Cincinnatian (Ordovician, Katian) small shelly phosphatic faunas dwarfed? Palaios 31, 55-70. View article

Davies, N.S. & Sansom, I.J. 2009. Ordovician vertebrate habitats: A Gondwanan perspective. Palaios 24, 717-722. View article

Devaere, L., Clausen, S., Álvaro, J.J., Peel, J.S. & Vachard, D. 2014. Terreneuvian orthothecid (Hyolitha) digestive tracts from northern Montagne Noire, France; taphonomic, ontogenetic and phylogenetic implications. PLoS One 9(2), e88583. View article

Di Cunzolo, S., Acenolaza, G.F. & Rodríguez Brizuela, R. 2003. Cruziana-Skolithos ichnoassociation in the Casa Colorada Formation (Upper Cambrian-Tremadocian), Cordillera Oriental of Jujuy Province, 285-288. In Albanesi, G.L., Beresi, M.S. & Peralta, S.H. (eds) Ordovician fron the Andes. Serie Correlación Geológica 17.

Donovan, S.K., den Ouden, N., Jagt, J.W.M. 2018. Preservation of borings: contrasting examples from the type Maastrichtian (Upper Cretaceous), southern Limburg, the Netherlands. Proceedings of the Geologists’ Association 129, 12-16. View article

Duperron, M., Scasso, R.A. & Moya, M.C. 2018. Geología del área del embalse Las Maderas, provincia de Jujuy, con referencia a las acumulaciones bioclásticas fosfáticas del Tremadociano y Floiano. Revista de la Asociación Geológica Argentina 75(1), 95-114.

Freeman, R.L., Dattilo, B.F. & Brett, C.E. 2019. An integrated stratinomic model for the genesis and concentration of “small shelly fossil”-style phosphatic microsteinkerns in not-so-exceptional conditions. Palaeogeography, Palaeoclimatology, Palaeoecology 535, 109344. View article

Gilmore, B. 1992. Scroll coprolites from the Silurian of Ireland and the feeding of early vertebrates. Palaeontology 35, 319-333.

Harper, D.A.T., Cascales-Minana, B. & Servais, T. 2020. Early Palaeozoic diversifications

and extinctions in the marine biosphere: a continuum of change. Geological Magazine 157, 5-21. View article

Hawkins, A.D., Liu, H.P., Briggs, D.E.G., Muscente, A.D., Mckay, R.M., Witzke, B.J. & Xiao, S. 2018. Taphonomy and biological affinity of three-dimensionally phosphatized bromalites from the Middle Ordovician Winneshiek Lagerstätte, northeastern Iowa, USA. Palaios 33, 1-15. View article

Henderson, R.A. & McNamara, K.J. 1985. Taphonomy and ichnology of cephalopod shells in a Maastrichtian chalk from Western Australia. Lethaia 18, 305-322. View article

Hunt, A.P. 1992. Late Pennsylvanian coprolites from the Kinney Brick Quarry, central New Mexico, with notes on the classification and utility of coprolites, 221-229. In Zidek, J. (ed.) Geology and paleontology of the Kinney Brick Quarry, Late Pennsylvanian, central New Mexico. New Mexico Bureau of Mines and Mineral Resources, Bulletin 138.

Hunt, A.P. & Lucas, S.G. 2012. Descriptive terminology of coprolites and recent feces, 153-160. In Hunt, A.P., Milán, J., Lucas, S.G. & Spielmann, J.A. (eds) Vertebrates Coprolites. New Mexico Museum of Natural History & Science, Bulletin 57.

Hunt, A.P. & Lucas, S.G. 2018. The record of sloth coprolites in North and South America: Implications for terminal Pleistocene extinctions. New Mexico Museum of Natural History & Science, Bulletin 79, 277-298.

Hunt, A.P. & Lucas, S.G. 2021. The Ichnology of Vertebrate Consumption. New Mexico Museum of Natural History and Science, Bulletin 87, 1-216.

Jeppsson, L., Anehus, R. & Fredholm, D. 1999. The optimal acetate buffered acetic acid technique for extracting phosphatic fossils. Journal of Paleontology 73(5), 964-972. View article

Knaust, D. 2008. Balanoglossites Magdefrau, 1932 from the Middle Triassic of Germany: part of a complex trace fossil probably produced by burrowing and boring polychaetes. Paläontologische Zeitschrift 82(4), 347-372. View article

Knaust, D. 2020a. Invertebrate coprolites and cololites revised. Papers in Palaeontology 6(3), 385-423. View article

Knaust, D. 2020b. Rhizocorallites Müller, 1955 from the Triassic and Jurassic of Germany: burrow, coprolite, or cololite? Paläontologische Zeitschrift 94, 769-785. View article

Lan, T., Yang, J., Hou, J.-B. & Zhang, X.-G. 2015. The feeding behaviour of the Cambrian tubiculous priapulid Selkirkia. Lethaia 48, 125-132. View article

Lucas, S.G. & Foley, M.P. 2023. Additions to the fossil biota of the Upper Cretaceous (Cenomanian) Paguate Member of the Dakota Formation, southeastern San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin 94, 297-300.

Maas, A., Huang, D., Chen, J., Waloszek, D. & Braun, A. 2007. Maotianshan-Shale nemathelminths - Morphology, biology, and the phylogeny of Nemathelminthes. Palaeogeography, Palaeoclimatology, Palaeoecology 254, 288-306. View article

Mángano, M.G. & Buatois, L.A. 2011. Timing of infaunalization in shallow-marine Early Paleozoic communities in Gondwanan settings: Discriminating evolutionary and paleogeographic controls. Palaeontologia Electronica 14(2), 1-21.

Mángano, M.G., Buatois, L.A. & Muniz Guinea, F. 2005. Ichnology of the Alfarcito Member (Santa Rosita Formation) of northwestern Argentina: animal-substrate interactions in a lower Paleozoic wave-dominated shallow sea. Ameghiniana 42(4), 641-668.

Mángano, M.G., Buatois, L.A., Waisfeld, B.G., Munoz, D.F., Vaccari, N.E. & Astini, R.A. 2021. Were all trilobites fully marine? Trilobite expansion into brackish water during the early Palaeozoic. Proceedings of the Royal Society B 288, 20202263. View article

Martin, E.L.O., Lerosey-Aubril, R. & Vannier, J. 2016. Palaeoscolecid worms from the Lower Ordovician Fezouata Lagerstätte, Morocco: Palaeoecological and palaeogeographical implications: Palaeogeography, Palaeoclimatology, Palaeoecology 460, 130-141. View article

Mastandrea, O. & Leanza, H.A. 1975. Los estratos con Lingula del norte argentino y su importancia como fuente de fosfatos. Congreso Iberoamericano de Geología Económica, Buenos Aires 2, 445-460.

Mayer, G. 1952. Neue Lebensspuren aus dem Unteren Hauptmuschelkalk (Trochitenkalk) von Wiesloch: Coprolus oblongus n. sp. und C. sphaeroideus n. sp. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1952, 376-379.

Müller, A.H. 1955. “Helminthoide” Lebensspuren aus der Trias von Thüringen. Geologie 4, 407-415.

Olempska, E., Błażejowski, B., Waloszek, D. & Maas, A. 2023. Phosphatic bromalites and microfossils from the Furongian (Cambrian) of northern Poland (Baltica) and palaeobiological implications. Palaeogeography, Palaeoclimatology, Palaeoecology 610, 111350. View article

Peel, J.S. 2015. Bromalites from the Cambrian (Series 2 and 3) of North Greenland. GFF 137, 181-194. View article

Reich, M. 2001. Ordovician holothurians from the Baltic Sea area, 93-96. In Barker, M. (ed.) Echinoderms 2000: Proceedings of the 10th International Conference, Dunedin. Balkema Publishers, Lisse.

Schalamuk, I.B., Fernández, R. & Etcheverry, R. 1983. Los Yacimientos de minerales no metalíferos y rocas de aplicación de la región NOA (Provincias de Catamarca, Jujuy, La Rioja, Salta y Tucumán). 205 pp. Ministerio de Economía, Subsecretaría de Minería, Buenos Aires.

Schallreuter, R. 1977. Eine neue fragliche kalkschalige Foraminifere aus einem mittelordovizischen Geschiebe Norddeutschlands. Paläontologische Zeitschrift 51(3-4), 227-233. View article

Schallreuter, R. 2003. New coprolites from the Ordovician of Thuringia and Siberia. Archiv für Geschiebekunde 4, 233-240.

Schweigert, G. 2001. Über Medusites Germar (Koprolith) und echte Algen aus oberjurassischen Plattenkalken Süddeutschlands. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2001, 237-249. View article

Servais, T., Cascales-Minana, B., Harper, D.A.T., Lefebvre, B., Munnecke, A., Wang, W. & Zhang, Y. 2023. No (Cambrian) explosion and no (Ordovician) event: A single long-term radiation in the early Palaeozoic: Palaeogeography, Palaeoclimatology, Palaeoecology 623, 111592. View article

Shen, C., Pratt, B.R. & Zhang, X.G. 2014. Phosphatized coprolites from the middle Cambrian (Stage 5) Duyun fauna of China. Palaeogeography, Palaeoclimatology, Palaeoecology 410, 104-112. View article

Stone, J. 1987. Review of investigative techniques used in the study of conodonts, 7-34. In Austin, R.L. (ed.) Conodonts: Investigative techniques and applications. Ellis Horwood Limited, Chichester.

Such, P., Buatois, L.A. & Mángano, M.G. 2007. Stratigraphy, depositional environments and ichnology of the Lower Paleozoic in the Azul Pampa area - Jujuy Province. Revista de la Asociación Geológica Argentina 62(3), 331-344.

Sumner, D. 1993. Coprolites from the Viséan of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Scotland, Earth Sciences 84, 413-416. View article

Tolmacheva, T.Y. & Purnell, M.A. 2002. Apparatus composition, growth, and survivorship of the Lower Ordovician conodont Paracordylodus gracilis Lindström, 1955. Palaeontology 45(2), 209-228. View article

Toom, U., Vinn, O., Isakar, M., Madison, A. & Hints, O. 2020. Small faecal pellets in Ordovician shelly fossils from Estonia, Baltoscandia. Estonian Journal of Earth Sciences 69, 1-19. View article

Toro, B.A., Arcerito, F.R.M., Munoz, D.F., Waisfeld, B.G. & De La Puente, G.S. 2015. Graptolite-Trilobite Biostratigraphy in the Santa Victoria Area, Northwestern Argentina. A Key for Regional and Worldwide Correlation of the Lower Ordovician (Tremadocian-Floian). Ameghiniana 52, 535-557. View article

Turner, J.C.M. 1960. Estratigrafía de la Sierra de Santa Victoria y adyacencias. Boletín de la Academia Nacional de Ciencias (Córdoba) 41(2), 163-196.

Uchman, A. & Rattazzi, B. 2023. Cork screw-like horizontal trace fossils with a focus on a new ichnospecies of Helicodromites from the Oligocene Molare Formation of NW Italy. Geological Quarterly 67, 22. View article

Vaucher, R., Vaccari, N.E., Balseiro, D., Munoz, D.F., Dillinger, A., Waisfeld, B.G. & Buatois, L.A. 2020. Tectonic controls on late Cambrian-Early Ordovician deposition in Cordillera oriental (Northwest Argentina). International Journal of Earth Sciences 109, 1897-1920. View article

Voldman, G.G. & Vaccari, N.E. 2025. Biostratigraphic implications of Kallidontus nodosus Pyle and Barnes in the Ordovician of the Eastern Cordillera, Argentina. Andean Geology 52, 203-220. View article

Voldman, G.G., Albanesi, G.L., Zeballo, F.J. & Monaldi, C.R. 2013a. Early Ordovician (late Floian) conodonts from the Zenta Range, Cordillera oriental, NW Argentina, 123-128. In Albanesi, G.L. & Ortega, G. (eds) Conodonts from the Andes, Proceedings of the 3th International Conodont Symposium, Asociación Paleontológica Argentina, Publicación Especial 13, Buenos Aires.

Voldman, G.G., Albanesi, G.L., Monaldi, C.R. & Zeballo, F.J. 2013b. An Early Ordovician conodont fauna from the Santa Rosita Formation at its type area in the Santa Victoria Range, Cordillera Oriental, Northwestern Argentina. Stratigraphy 10(4), 301-322. View article

Voldman, G.G., Albanesi, G.L., Ortega, G., Giuliano, M.E. & Monaldi, C.R. 2017. New conodont taxa and biozones from the Lower Ordovician of the Cordillera Oriental, NW Argentina. Geological Journal 52, 394-414. View article

Waisfeld, B.G. & Vaccari, N.E. 2003. Trilobites, 295-410. In Benedetto, J.L. (ed.) Ordovician Fossils of Argentina. Universidad Nacional de Córdoba, Secretaría de Ciencia y Tecnología, Córdoba.

Waisfeld, B.G., Benedetto, J.L., Toro, B.A., Voldman, G.G., Rubinstein, C.V., Heredia, S., Assine, M.L., Vaccari, N.E. & Niemeyer, H. 2023. The Ordovician of southern South America, 133-173. In Servais, T., Harper, D.A.T., Lefebvre, B., Percival, I.G. (eds) A Global Synthesis of the Ordovician System: Part 2. Geological Society London, Special Publication 532. View article

Waterson, C.D., Oelofsen, B.N. & Oostuizen, R.D.F. 1985. Cyrtoctenus wittbergensis sp. nov. (Chelicerata: Eurypterida) a large sweep-feeder from the Carbonifeous of South Africa. Transactions of the Royal Society of Scotland, Earth Sciences 76, 339-358. View article

Zeballo, F.J. 2011. Bioestratigrafía de conodontes y graptolitos, y fósiles asociados, de las secuencias cambro-ordovícicas del flanco oriental de la Quebrada de Humahuaca, provincia de Jujuy, Argentina. 362 pp. Ph.D. thesis, Universidad Nacional de Córdoba, Córdoba, Argentina.

Zeballo, F.J. & Albanesi, G.L. 2013. New conodont species and biostratigraphy of the Santa Rosita Formation (upper Furongian-Tremadocian) in the Tilcara Range, Cordillera Oriental of Jujuy, Argentina. Geological Journal 48, 170-193. View article

Zeballo, F.J., Albanesi, G.L., Voldman, G.G. & Monaldi, C.R. 2013. New records of Tremadocian conodonts from the Zenta Range, Jujuy Province, Argentina. Conodonts from the Andes, Asociación Paleontológica Argentina, publicación especial 13, 129-133.

Zhen, Y.Y. 2019. Conodont biostratigraphy of the Horn Valley Siltstone, Amadeus Basin in central Australia, 253-255. In 13th International Symposium on the Ordovician System: Contributions of International Symposium. Novosibirsk, Russia (July 19-22, 2019), Publishing House of SB RAS, Novosibirsk.

Zhen, Y.Y. & Nicoll, R.S. 2009. Biogeographic and biostratigraphic implications of the Serratognathus bilobatus fauna (Conodonta) from the Emanuel Formation (Early Ordovician) of the Canning Basin, Western Australia. Records of the Australian Museum 61(1), 1-30. View article