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Who ate whom? Paleoecology revealed through synchrotron microtomography of coprolites (fossil feces)
Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Organismal Biology.ORCID iD: 0000-0001-7998-2243
2020 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

Fossil droppings, known as coprolites, are being increasingly recognized as a valuable source of paleoecological information with special regard to diets, parasitism, and physiology of extinct taxa. Here, it is suggested that the excellent preservation and amount of inclusions in the coprolites (e.g. food residues and parasites) qualifies them as Lagerstätten – deposits with exceptional paleontological information. However, two interlinked problems commonly arise when they are studied. Firstly, it is often difficult to tie coprolites to producers and, secondly, it is challenging to recognize the fragmented and randomly distributed inclusions in their matrix. Here I use propagation phase-contrast synchrotron microtomography (PPC-SRμCT) in combination with other techniques to solve these problems. As a result, the oldest known example of archosaurian osteophagy is uncovered based on inter alia the occurrence of serrated teeth and many crushed bones in coprolites assigned to the Late Triassic theropod-like archosaur Smok wawelski. Osteophagy has previously been thought to be rare among extinct archosaurs with the exception of Late Cretaceous tyrannosaurids. This suggests some degree of ecological convergence between the tyrannosaurids and S. wawelski. Furthermore, exceptionally-preserved beetle remains are discovered in coprolites tentatively assigned to the Triassic dinosauriform Silesaurus opolensis, which had a specialized dentition and possessed beak-shaped jaws that were likely used to peck insects off the ground. Moreover, pterosaur coprolites are shown to contain similar food residues as found in droppings of recent flamingos, implying that some Late Jurassic pterosaurs were filter feeders. I argue that such paleoecological studies have a large impact on our understanding of ancient animals, and that studies of coprolites can unravel parts of ancient food webs in unprecedented ways. Information on past food webs may, in turn, be used to analyze trophic changes through time, which could cast new light on big evolutionary events. This is demonstrated by reconstructing trophic structures in early Mesozoic assemblages that represent snapshots of three stages of early dinosaur evolution.

Place, publisher, year, edition, pages
Uppsala: Acta Universitatis Upsaliensis, 2020. , p. 49
Series
Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology, ISSN 1651-6214 ; 1904
Keywords [en]
coprolites, paleoecology, synchrotron microtomography, taphonomy, Triassic
National Category
Other Biological Topics
Research subject
Biology
Identifiers
URN: urn:nbn:se:uu:diva-404162ISBN: 978-91-513-0875-3 (print)OAI: oai:DiVA.org:uu-404162DiVA, id: diva2:1393402
Public defence
2020-04-03, Lindahlsalen, Evolutionsbiologiskt centrum, Norbyvägen 14, Uppsala, 09:15 (English)
Opponent
Supervisors
Available from: 2020-03-13 Created: 2020-02-16 Last updated: 2020-05-19
List of papers
1. Vertebrate coprolites (fossil faeces): An underexplored Konservat-Lagerstatte
Open this publication in new window or tab >>Vertebrate coprolites (fossil faeces): An underexplored Konservat-Lagerstatte
2016 (English)In: Earth-Science Reviews, ISSN 0012-8252, E-ISSN 1872-6828, Vol. 162, p. 44-57Article, review/survey (Refereed) Published
Abstract [en]

Fossilized soft tissues of animals (e.g. muscles, hair and feathers) are valuable sources of palaeobiological information, but a poor preservation potential makes them undesirably scarce in the fossil record. The aim of this review is to summarize main findings, current progress and the analytical constraints of detecting fossilized soft tissues in coprolites from, mainly, freshwater and terrestrial carnivorous vertebrates. We conclude that soft-tissue inclusions in coprolites are sources of two important lines of information: the fossils can be put in a direct palaeoecological context, and characters of extinct taxa are more likely preserved in the phosphate-rich taphonomic microenvironment of coprolites than elsewhere. As a result, it is possible to unravel the deep-time origins of host-parasite relations, to understand ancient trophic food webs and detect new soft-tissue characters of different animal groups. Examples of the latter include muscle tissues from a tyrannosaurid prey, tapeworm eggs (including a developing embryo) in a Permian shark coprolite, as well as hair from multituberculates and, probably, from stem-mammals (Therapsids). Additionally, the use of coprolites in an archaeological context is briefly reviewed with focus on key aspects that may become implemented in studies of pre-Quaternary specimens as well. In summary, there is a wide range of information that can be extracted from coprolites, which has not yet been fully explored in palaeontological studies.

Keywords
Coprolites, Soft tissues, Lagerstatten, Phosphatization, Conservation traps, Palaeoecology
National Category
Geosciences, Multidisciplinary Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-311769 (URN)10.1016/j.earscirev.2016.08.014 (DOI)000388776700003 ()
Funder
Knut and Alice Wallenberg FoundationSwedish Research Council, 2014-4367
Available from: 2017-01-02 Created: 2017-01-02 Last updated: 2020-02-16Bibliographically approved
2. Synchrotron phase-contrast microtomography of coprolites generates novel palaeobiological data
Open this publication in new window or tab >>Synchrotron phase-contrast microtomography of coprolites generates novel palaeobiological data
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2017 (English)In: Scientific Reports, E-ISSN 2045-2322, Vol. 7, article id 2723Article in journal (Refereed) Published
Abstract [en]

Coprolites (fossil faeces) reveal clues to ancient trophic relations, and contain inclusions representing organisms that are rarely preserved elsewhere. However, much information is lost by classical techniques of investigation, which cannot find and image the inclusions in an adequate manner. We demonstrate that propagation phase-contrast synchrotron microtomography (PPC-SR mu CT) permits high-quality virtual 3D-reconstruction of coprolite inclusions, exemplified by two coprolites from the Upper Triassic locality Krasiejow, Poland; one of the coprolites contains delicate beetle remains, and the other one a partly articulated fish and fragments of bivalves.

Place, publisher, year, edition, pages
Nature Publishing Group, 2017
National Category
Natural Sciences
Identifiers
urn:nbn:se:uu:diva-327451 (URN)10.1038/s41598-017-02893-9 (DOI)000402515800041 ()28578409 (PubMedID)
Available from: 2017-08-11 Created: 2017-08-11 Last updated: 2022-09-15Bibliographically approved
3. Tyrannosaurid-like osteophagy by a Triassic archosaur
Open this publication in new window or tab >>Tyrannosaurid-like osteophagy by a Triassic archosaur
2019 (English)In: Scientific Reports, E-ISSN 2045-2322, Vol. 9, article id 925Article in journal (Refereed) Published
Abstract [en]

Here we present evidence for osteophagy in the Late Triassic archosaur Smok wawelski Niedzwiedzki, Sulej and Dzik, 2012, a large theropod-like predator from Poland. Ten medium to large-sized coprolites are matched, by their dimensions and by association with body fossils and footprints, to S. wawelski. The coprolites contain fragments of large serrated teeth as well as up to 50 percent by volume of bone fragments, with distinct fragmentation and angularity, from several prey taxa. This suggests pronounced osteophagy. Further evidence for bone-crushing behaviour is provided by isolated worn teeth, bone-rich regurgitalites (fossil regurgitates) and numerous examples of crushed or bite-marked dicynodont bones, all collected from the same bone-bearing beds in the Lipie Slaskie clay-pit. Several of the anatomical characters related to osteophagy, such as a massive head and robust body, seem to be shared by S. wawelski and the tyrannosaurids, despite their wide phylogenetic separation. These large predators thus provide evidence of convergence driven by similar feeding ecology at the beginning and end of the age of dinosaurs.

National Category
Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-377342 (URN)10.1038/s41598-018-37540-4 (DOI)000457128700002 ()30700743 (PubMedID)
Funder
Swedish Research Council, 2017-05248Knut and Alice Wallenberg Foundation
Available from: 2019-02-25 Created: 2019-02-25 Last updated: 2022-09-15Bibliographically approved
4. Beetle-bearing coprolites possibly reveal the diet of a Late Triassic dinosauriform
Open this publication in new window or tab >>Beetle-bearing coprolites possibly reveal the diet of a Late Triassic dinosauriform
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2019 (English)In: Royal Society Open Science, E-ISSN 2054-5703, Vol. 6, no 3, article id 181042Article in journal (Refereed) Published
Abstract [en]

Diets of extinct animals can be difficult to analyse if no direct evidence, such as gut contents, is preserved in association with body fossils. Inclusions from coprolites (fossil faeces), however, may also reflect the diet of the host animal and become especially informative if the coprolite producer link can be established. Here we describe, based on propagation phase-contrast synchrotron microtomography (PPC-SRμCT), the contents of five morphologically similar coprolites collected from two fossil-bearing intervals from the highly fossiliferous Upper Triassic locality at Krasiejow in Silesia, Poland. Beetle remains, mostly elytra, and unidentified exoskeleton fragments of arthropods are the most conspicuous inclusions found in the coprolites. The abundance of these inclusions suggests that the coprolite producer deliberately targeted beetles and similar small terrestrial invertebrates as prey, but the relatively large size of the coprolites shows that it was not itself a small animal. The best candidate from the body fossil record of the locality is the dinosauriform Silesaurus opolensis Dzik, 2003, which had an anatomy in several ways similar to those of bird-like neotheropod dinosaurs and modern birds. We hypothesize that the beak-like jaws of S. opolensis were used to efficiently peck small insects off the ground, a feeding behaviour analogous to some extant birds.

Keywords
coprolites, palaeoecology, Dinosauromorpha, beetle elytra, insectivore
National Category
Geosciences, Multidisciplinary Other Earth and Related Environmental Sciences
Identifiers
urn:nbn:se:uu:diva-383880 (URN)10.1098/rsos.181042 (DOI)000465470300008 ()31031991 (PubMedID)
Funder
Swedish Research Council, 2017-05248
Available from: 2019-06-11 Created: 2019-06-11 Last updated: 2020-02-16Bibliographically approved
5. Filter feeding in Late Jurassic pterosaurs supported by coprolite contents
Open this publication in new window or tab >>Filter feeding in Late Jurassic pterosaurs supported by coprolite contents
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2019 (English)In: PeerJ, E-ISSN 2167-8359, Vol. 7, article id e7375Article in journal (Refereed) Published
Abstract [en]

Diets of pterosaurs have mainly been inferred from indirect evidence such as comparative anatomy, associations of co-occurring fossils, and functional morphology. Gut contents are rare, and until now there is only a single coprolite (fossil dropping), with unidentified inclusions, known. Here we describe three coprolites collected from a palaeosurface with numerous pterosaur tracks found in early Kimmeridgian (Hypselocyclum Zone) intertidal deposits of the Wierzbica Quarry, Poland. The specimens' morphology and association to the tracks suggest a pterosaur producer. Synchrotron scans reveal numerous small inclusions, with foraminifera making up the majority of the identifiable ones. Other small remains include shells/carapaces (of bivalves, ostracods, and other crustaceans/arthropods) and bristles (some possibly of polychaete worms). The high density of the small shelly inclusions suggest that they were not accidently ingested, but constituted an important food source for the pterosaur(s), perhaps together with unpreserved soft-bodied animals. The combined evidence from the tracks and coprolites suggest a filter-feeding ctenochasmatid as the most likely tracemaker. If true, this significantly expands the bromalite record for this pterosaur group, which was previously only known from gastroliths. Moreover, this study also provides the first direct evidence of filter feeding in Jurassic pterosaurs and shows that they had a similar diet to the recent Chilean flamingo (Phoenicopterus chilensis).

Place, publisher, year, edition, pages
PEERJ INC, 2019
Keywords
Filter feeding, Coprolites, Pterosaur, Palaeoecology, Ctenochasmatidae, Late Jurassic
National Category
Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-398849 (URN)10.7717/peerj.7375 (DOI)000482716000001 ()31523493 (PubMedID)
Funder
Swedish Research Council, 2017-05248
Available from: 2019-12-19 Created: 2019-12-19 Last updated: 2023-08-28Bibliographically approved
6. Food web analysis across the Triassic-Jurassic transition shed new light on early dinosaur evolution and herbivore diversity
Open this publication in new window or tab >>Food web analysis across the Triassic-Jurassic transition shed new light on early dinosaur evolution and herbivore diversity
Show others...
(English)Manuscript (preprint) (Other academic)
Keywords
palaeoecology, coprolites, trophic structure, dinosaurs, Triassic, Jurassic
National Category
Biological Sciences
Identifiers
urn:nbn:se:uu:diva-404254 (URN)
Available from: 2020-02-16 Created: 2020-02-16 Last updated: 2020-02-17Bibliographically approved

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