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Evolution and Taxonomy of Cambrian Arthropods from Greenland and Sweden

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Evolution and Taxonomy of Cambrian Arthropods from Greenland and Sweden Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 558 Evolution and taxonomy of Cambrian arthropods from Greenland and Sweden MARTIN STEIN ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 UPPSALA ISBN 978-91-554-7302-0 2008 urn:nbn:se:uu:diva-9301 !! "#!$% & ' $( !))* ()+)) , - . , / , !))*, 0 1 /. , 2 , ##*, $# , , 3/4 5"*65(6##76"$)!6), 2 8 , 4 8 , & . 8 , 0 1 . 0 / 9 , - . 0 / ! 1 , - . / 9 :' : 0 /. , ; 1 / ! $ 1 , 3 /<. & =< - . 9 , 2 . 3 & 6 1 . , . , ; 1 & , 3 . 6 2 :/ & 6> / . ! 4 & . 3 & , ?' @ 0 $ 7 /. 0 , " # . / 9 . < : : , - : : $ . $ 0 . , & . . ? @ < , " ! % 0 2 - 1 /. & ' ( ' )* +,' ' (-./01, ' ! A / !))* 3// (%#(6%!(7 3/4 5"*65(6##76"$)!6) + +++ 65$)( +BB ,<,B C D + +++ 65$)( List of publications I. Stein, M. 2008. Fritzolenellus lapworthi (Peach and Horne, 1892) from the lower Cambrian (Cambrian Series 2) Bastion Formation of North-East Greenland. Bulletin of the Geological Society of Denmark 56:1–10. II. Stein, M. & Peel, J. S. 2008. Perissopyge (Trilobita) from the lower Cambrian (Series 2, Stage 4) of North America and Greenland. GFF 130:71–78. III. Stein, M., Waloszek, D., Maas, A., Haug, J. T. & Müller, K. J. 2008. The stem crustacean Oelandocaris oelandica re-visited. Acta Palaeon- tologica Polonica 53:461–484. IV. Stein, M. (manuscript) A new arthropod from the Early Cambrian of North Greenland with a ‘great appendage’ like antennula. Submitted to Zoological Journal of the Linnean Society. V. Stein, M., Peel, J. S., Siveter, D. J. & Williams, M. (manuscript) Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte, lower Cambrian of North Greenland. VI. Lagebro, L., Stein, M. & Peel, J. S. (manuscript) A new arthropod from the early Cambrian Sirius Passet Fauna of North Greenland. Reproduction of the published papers is made with kind permission from the copyright holders: Paper I © Bulletin of the Geological Society of Denmark, Dansk Geologisk Forening Paper II © GFF, Geologiska Föreningen Paper III © Acta Palaeontologica Polonica, Instytut Paleobiologii PAN Statement of authorship Access to the material studied in papers I,–II, IV–VI was granted by J. S. Peel. Paper I: M. Stein performed the studies and wrote the manuscript. Paper II: M. Stein carried out the taxonomic studies and wrote the section on systematic palaeontology. J. S. Peel provided the introduction and the sec- tions on material and stratigraphy. Paper III: M. Stein and D. Waloszek carried out most of the studies of the material. Initial reconstructions were created by M. Stein, D. Waloszek, and A. Maas. M. Stein created an early version of the three dimensional model of the largest stage and the animated sequence; J. T. Haug created the final ver- sion of the model of the largest stage and all models of the ontogenetic se- quence. K. J. Müller collected the material. Manuscript mainly by M. Stein and D. Waloszek with contributions by the other authors. Paper IV: M. Stein performed the studies, created the reconstructions and wrote the manuscript. Paper V: Initial studies were carried out by M. Stein and J. S. Peel. Detailed studies were carried out by M. Stein, D. Siveter and M. Williams. Manuscript mainly by M. Stein with contributions by the other authors. Paper VI: L. Lagebro provided an initial study of the material. M. Stein per- formed detailed restudy and wrote the section on systematic palaeontology. J. S. Peel wrote the introduction. Disclaimer The papers presented here are for the purpose of public examination as a doctoral thesis only. They are not considered publications according to the ICZN. Accordingly, all new taxonomic names and emendations in papers IV–VI are void. Authority for taxonomic work in papers I–III is by the origi- nal publications. Front cover Reconstruction of Kiisortoqia avannaarsuensis from the early Cambrian Sir- ius Passet Lagerstätte of Greenland. Contents Introduction ................................................................................................... 7 Trilobite biostratigraphy ................................................................................ 9 Paper I ..................................................................................................... 10 Paper II .................................................................................................... 12 Arthropod phylogeny ................................................................................... 14 The current status of euarthropod phylogeny from a neontological per- spective ................................................................................................... 15 The ground pattern and the importance of fossils ................................... 16 The position of Trilobita ......................................................................... 19 Paper III .................................................................................................. 21 Paper IV .................................................................................................. 22 Paper V .................................................................................................... 24 Paper VI .................................................................................................. 25 Svensk sammanfattning ............................................................................... 27 Trilobit-stratigrafi på Grönland ............................................................... 28 De tidiga leddjurens evolution ................................................................ 29 Acknowledgments ....................................................................................... 30 References ................................................................................................... 31 Introduction Arthropods are a diverse group of multicellular animals (metazoans), esti- mated to account for more than three quarters of the present-day metazoan diversity. They constitute an important element of many ecosystems, both marine and terrestrial. Four major groups are recognized among extant arthropods. The chelicerates include the familiar spiders, scorpions, mites, and ticks, and less familiar forms such as the horseshoe crabs. Crabs and lob- sters are grouped together as crustaceans, along with woodlice and krill, but also with smaller and less familiar forms such as brine-shrimps and the truly ubiquitous ostracodes and copepods, and even sessile forms such as barna- cles. Millipedes and centipedes comprise the myriapods, but the all but too familiar insects form the largest of all living arthropod groups. These four groups sharing a common ancestor constitute the Euarthropoda and are char- acterized by a unique body structure and movement apparatus where mus- cles do not act as antagonists to bones or fluid-filled pressure vessels lodged inside a ‘soft’ body. Rather, a firm cuticle, which is produced by the epider- mal cells surrounding the body, forms an ‘exoskeleton’ consisting of hard- ened (sclerotized) plates conjoined by soft membranous regions that allow for flexibility. Muscles, attached to the sclerotized parts, act against this ‘ex- oskeleton’. How the euarthropod groups are related to each other remains the subject of sometimes contentious debate. The nature of several small groups considered to be the closest relatives to the euarthropods is also debated. While there is consensus that the little known onychophorans (velvet worms) are arthropods in the wider sense, researchers still argue about if the minute tardigrades (water bears) belong to the group or are more closely related to roundworms. The pentastomids (tongue worms), a group of worm-like para- sites infesting the respiratory organs of land-living vertebrates (amphibians, ‘reptiles’, birds, and mammals) are widely accepted as arthropods. But are they the sister group to euarthropods or highly derived crustaceans, and thus euarthropods? Given their tremendous diversity, it is not surprising that arthropods have had a long evolutionary history, evident not least in a rich fossil record span- ning the whole Phanerozoic. Most common in the fossil record are forms with well biomineralized (calcified) ‘exoskeletons’, such as trilobites or cer- tain crustaceans, including ostracodes and malacostracans. Trilobites ap- peared early in the Cambrian and flourished through the whole Palaeozoic 7 prior to disappearing in the terminal Permian extinction. They are common and diverse components of Cambrian faunas and have been of paramount importance in biostratigraphic and palaeogeographic studies of that period for more than a century and a half. But the fossil record of arthropods is not just limited to forms with calci- fied ‘exoskeletons’. Throughout the Phanerozoic, examples of exceptional preservation of soft parts offer unique glimpses into past ecosystems. Arthro- pods form a surprisingly common element of these so-called lagerstätten, al- though taphonomic bias against various
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