The Morphology and Evolutionary Significance of the Anomalocaridids - DocsLib

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Home , Amplectobelua, Buenellus, Hurdia, Tamisiocaris

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List of Papers

This thesis is based on the following papers, which are referred to in the text by their Roman numerals.

I Daley, A.C., Budd, G.E., Caron, J.-B., Edgecombe, G.D. & Collins, D. 2009. The Burgess Shale anomalocaridid Hurdia and its significance for early euarthropod evolution. Science, 323:1597-1600. II Daley, A.C. & Budd, G.E. New anomalocaridid appendages from the Burgess Shale, Canada. In press. Palaeontology. III Daley, A.C., Budd, G.E. and Caron, J.-B. The morphology and systematics of the anomalocaridid Hurdia from the Burgess Shale. Manuscript to be submitted to Journal of Systematic Pa- leontology. IV Daley, A.C. & Peel, J.S. A possible anomalocaridid from the Cambrian Sirius Passet Lagerstätte, North Greenland. In press. Journal of Paleontology. V Budd, G.E. & Daley, A.C. The lobes and lobopods of Opabinia regalis. Submitted to Lethaia.

Additionally, the following papers were prepared during the course of my PhD studies, but are not included in the thesis:

VI Daley, A.C. 2008. Statistical analysis of mixed-motive shell borings in Ordovician, Silurian and Devonian brachiopods from northern and eastern Canada. Canadian Journal of Earth Sci- ences, 45:213-229. VII Fischer, A.H.L., Arboleda, E., Egger, B., Hilbrant, M., McGregor, A.P., Cole, A.G., and Daley, A.C. 2009. ZOONET: Perspectives on the Evolution of Animal Form. Meeting Report. Journal of Experimental Zoology, Molecular and Developmen- tal Evolution, 312B:679-685. VIII Caron, J.-B., Gaines, R.R., Mángano, G.M., Streng, M. and Daley, A.C. A new Burgess Shale-type assemblage from the “thin” Stephen Formation of the Southern Canadian Rockies. Submitted to Geology.

Reprinting and publication is made with permission from the respective copyright holders.

Paper I, © The American Association for the Advancement of Science. Papers II, III, IV, V are copyright of the authors.

Statement of authorship

Paper I: A.C.D collected data, prepared figures, wrote the manuscript to- gether with G.E.B., and ran the phylogenetic analysis with G.D.E. and G.E.B. Fossil material and assemblages were collected, prepared and identified by D.C., and J.B.C. also prepared some specimens. All authors contributed to the interpretation of the fossils and commented on the manuscript.

Paper II: A.C.D. collected data, prepared figures, and wrote the manuscript. Both authors contributed to the interpretation of fossil material, and G.E.B commented on the manuscript.

Paper III: A.C.D. collected data, ran the statistical analyses, contributed to the interpretation, and prepared figures. All authors contributed to the interpretation of the fossils and wrote the manuscript.

Paper IV: A.C.D. prepared the figures and wrote the manuscript. J.S.P. collected, prepared and photographed the specimens. Both authors contributed to the fossil interpretation, and J.S.P. commented on the manuscript.

Paper V: A.C.D. prepared the figures. Both authors photographed the specimens, contributed to the fossil interpretation and wrote the manuscript.

Disclaimer

The papers presented here are for the purpose of public examination as a doctoral thesis only. They are not deemed valid for taxonomic or nomencla- tural purposes [see article 8.2 in the International Code of Zoological No- menclature 4th Edition, edited by Ride et al. (2000)]. Accordingly, all new taxonomic names and emendations in paper III are void. Authority for taxonomic work in papers I, II and IV is retained by the original publications.

Front cover: Reconstruction of Hurdia by Marianne Collins, 2008 ©ROM/J.-B. Caron. Used with permission.

Contents

Introduction ...... 9 The early evolution of animal life ...... 9 Euarthropod stem lineage ...... 11 The anomalocaridids ...... 14 History of description from the Burgess Shale...... 14 The Burgess Shale anomalocaridid Hurdia victoria ...... 18 Diversity and global distribution of the anomalocaridids ...... 20 Anomalocaridid diversity at the Burgess Shale ...... 20 A possible anomalocaridid from Sirius Passet, Greenland ...... 22 The lobes and lobopods of Opabinia ...... 24 Conclusions and future perspectives ...... 28 Svensk sammanfattning ...... 29 Hurdia victoria från Burgess Shale ...... 29 Diversitet och spridning av anomalocarider ...... 30 Morfologin hos Opabinia regalis ...... 31 Acknowledgements ...... 32 References ...... 34

Introduction

The early evolution of animal life The evolution of animal life extends back over half a billion years to a time when a major evolutionary radiation, the “Cambrian Explosion”, gave rise to nearly all of the major animal phyla known today (Conway Morris, 1989a; Budd, 2003). Occurring between about 600 and 500 million years ago, this radiation is exquisitely recorded by various fossil lagerstätten, where excep- tional preservation of soft-bodied animals provides a glimpse of complex marine ecosystems during the earliest evolution of modern animal life (Conway Morris, 1989b). Many Cambrian fossils are bizarre and enigmatic forms quite unlike their modern descendants, often inciting lively debate over their morphology, affinity and evolutionary significance. One such group of animals is the anomalocaridids, large presumed predators with a complicated history of description. The following thesis presents research on the morphology, systematics and evolutionary significance of the anomalocaridids and the closely related Opabinia regalis.

Most of the fossils studied herein come from the Burgess Shale in the Ca- nadian Rocky Mountains, the most famous of all Cambrian lagerstätten. Since its discovery by Charles D. Walcott in 1909, the Burgess Shale has produced thousands of exceptionally preserved fossils of soft-bodied organisms representing most major animal groups. The main sites are located on Fossil Ridge and Mount Stephen, near the small town of Field, British Co- lumbia (Fig. 1A). The Burgess Shale Formation is a series of basinal cal- careous argillaceous rocks of latest Glossopleura to early Bathyriscus- Elrathina zonal age that has been divided into 10 members (Fig. 1B), most of which have produced fossils (Fletcher and Collins, 1998). The exact pa- leoenvironment and mode of preservation of the Burgess Shale is controversial. Some theories suggest that organisms were living along the Cathedral Escarpment, and were transported to deeper water environments by mudflows (Conway Morris, 1979, 1986), while others argue that most organisms lived, died, and decayed in the same local environment where they were deposited and rapidly buried by mudflows (Caron and Jackson, 2006), possibly in close proximity to mud mounds and brine seeps in some localities other than those on Fossil Ridge (Collom et al., 2009; Johnson et al., 2009). The mode of preservation of Burgess Shale-type fossils involved the sup-

9 pression of organic degradation in the marine environment (Gaines et al., 2008), perhaps by rapid occlusion of sediment porosity soon after deposition (Gaines et al., 2005), or through complex chemical reactions in the sediment, such as clay-organic interactions (Butterfield, 1995) or adsorption of Fe3+ onto biopolymers (Petrovich, 2001). Early authigenic mineral replacement (Orr et al., 1998) and/or late diagenetic aluminosilicate mineral emplacement (Butterfield et al., 2007) also preserve certain anatomical features of some specimens. The main collections of Burgess Shale material include the original specimens collected by Walcott between the years 19091917, deposited at the Smithsonian National Museum of Natural History (USNM) in Wash- ington, a small collection at the Geological Survey of Canada (GSC) from quarrying activities in 1966 and 1967, and an extensive collection at the Royal Ontario Museum (ROM) from quarrying and prospecting activities beginning in 1975 and continuing to this day. Research presented in the following thesis is based mainly on material housed in these collections, as well as a small number of specimens from the Harvard University Museum of Comparative Zoology (MCZ) and the Yale Peabody Museum of Natural History (YPM).

Figure 1. A, Location map of the Burgess Shale and Sirius Passet fossil lagerstätten. B, Stratigraphy of the Burgess Shale Formation from Fletcher and Collins (1998) and Vannier et al. (2007).

A less well-known Cambrian fossil lagerstätte is the Sirius Passet biota in Peary Land, North Greenland (Fig. 1A). It was first described by Conway Morris et al. (1987), and consists of a series of black laminated mudstones and fine siltstones belonging to the Buen Formation, deposited adjacent to a buried carbonate escarpment (Babcock and Peel, 2007). It is 10 my older than the Burgess Shale, and is the oldest Cambrian fossil lagerstätte in Laurentia, possibly predating the Chengjiang fossil lagerstätte of southern China (Chen and Zhou, 1997; Hou et al., 2004; Conway Morris and Peel, in

10 press; Budd, in press). The fauna consists of mineralized and soft-bodied arthropods, and although large stem-group arthropods such as the lobopodians Kerygmachela and Pambdelurion have been described from this locality, anomalocaridids had not. One chapter of this thesis describes for the first time a possible anomalocaridid from this locality, based on a single isolated frontal appendage house at the Geological Museum (State Museum of Natu- ral History), Copenhagen, Denmark (MGUH).

Euarthropod stem lineage Arthropoda constitutes the largest and most diverse animal group on the planet today, with over three-quarters of known living and fossil species belonging to this invertebrate phylum. The arthropods are characterized by the possession of a chitino-proteinaceous exoskeleton formed of sclerotized plates joined by softer membranes, giving them jointed limbs and trunks. Four main groups of modern arthropods are recognized: the chelicerates, including spiders, mites, scorpions, ticks and horseshoe crabs; the myriapods, composed of centipedes and millipedes; the crustaceans, including lobsters, crabs, barnacles, ostracods and brine shrimp; and the largest arthropod group, the insects. Relationships between the main arthropod groups remain a subject of debate, and must also involve the consideration of other animal groups related to arthropods in a broad sense, including the onychophorans, or velvet worms, and possibly the tardigrades, or water bears. As proportionally abundant in the fossil record as they are in the modern world, the arthropods have a history that extends back to the early Cambrian, and are a major constituent of both normal mineralized fossil deposits and fossil lagerstätten. At the Burgess Shale (Conway Morris, 1986; Caron and Jackson, 2008) and Sirius Passet (Conway Morris et al., 1987; Babcock and Peel, 2007) soft-bodied arthropods and arthropod-like taxa dominate the biota and possess a diverse range of morphologies, some of them differing greatly from those of the four main groups of arthropods today. These unusual taxa are best considered in light of the stem and crown group concept (Budd and Jensen, 2000; Briggs and Fortey, 2005), which allows for the integration of fossil data, even as far back as the Cambrian, into modern animal phylogeny (Patterson, 1981). A crown group includes all extant (living) members of a monophyletic clade that has a single common ancestor, as well as all descendents of that ancestor (Fig. 2). Extinct (fossil) taxa that are more closely related to a crown group than any other crown group, but fall outside of it because they are missing key morphological characteristics that define the group, are considered as stem taxa (Budd and Jensen, 2000). Stem taxa form a paraphyletic lineage leading up to modern crown groups, providing us with a record of actual evolutionary history and giving insight into the order of character acquisition and possible homologies between living taxa.

11

Figure 2. Generalized phylogenetic tree of selected stem- and crown-group arthropods. Based on the cladistic analysis in Paper I. Illustrations by M. Streng.

The membership of the stem lineage of the arthropods is somewhat controversial, particularly in regards to certain features of key fossil taxa. Cam- brian fossil lagerstätten, in particular the Burgess Shale, Sirius Passet and Chengjiang biota, host a plethora of taxa with varying degrees of arthropodi- zation (Budd and Telford, 2009; Edgecombe, in press). Although the specif- ics are intensely debated, there are several major points of consensus about the arthropod stem lineage (Fig. 2), giving a view of general evolutionary trends. The basal part of the stem lineage is occupied by Onychophora and a grade of several taxa of Cambrian lobopodians, which have annulated (as opposed to segmented) bodies bearing unjointed lobopod walking limbs and a pair of unmineralized frontal appendages. Further uptree are the “gilled lobopods” from Sirius Passet, Kerygmachela (Budd, 1993) and Pambde- lurion (Budd, 1998), characterized by the possession of an unsclerotized frontal appendage pair, walking lobopods limbs and lateral swimming lobes with associated gill structures. These taxa are typically located either below (Budd, 1996, 1999, 2002; Dewel and Dewel, 1998; Cotton and Braddy, 2004; Kühl et al., 2009; Paper I) or in a clade with (Chen et al., 1994; Hou and Bergström, 2006; Ma et al., 2009) Opabinia and the anomalocaridids. Opabinia has a segmented body with lateral swimming lobes, and a cephalon bearing five eyes and a proboscis with spines at the end (Whittington, 1975). The presence of lobopod walking limbs in this animal is a highly

12 contentious issue (Budd, 1996; Zhang and Briggs, 2007) discussed in Paper V of this thesis. The Burgess Shale anomalocaridids also possess lateral swimming lobes, but no evidence of walking limbs, as well as a pair of cu- ticularized frontal appendages, a circular sclerotized mouth part, and a pair of eyes on stalks (Whittington and Briggs, 1985). Above the anomalocaridids, a grade of upper stem group arthropods includes a wide variety of taxa with unresolved relationships (Hou and Bergström, 1997; Budd, 2002, 2008; Waloszek et al., 2005, 2007; Bergström et al., 2008), but which have hardened tergites, biramous limbs with a segmented walking branch and an outer flap, and a “short” frontal appendage (e.g. Canadaspis, Fuxianhuia, and Leanchoilia). A full discussion of all the issues surrounding the arthropod stem lineage is beyond the scope of this thesis, however one topic that is elaborated upon is the origin and evolution of the arthropod biramous limb. Extant arthropod limbs vary widely in morphology, but are of two major types, either uniramous, as seen in the chelicerates (except for the horseshoe crabs), insects and myriapods, or biramous as seen in the crustaceans. Biramy is also widespread in upper stem group arthropods (e.g. Hou and Bergström, 1997; Hou et al., 2004), however taxa in the lower part of the stem lineage, such as the lobopodians and onychophorans, possess uniramous lobopod limbs. These ventrally directed limbs are considered to be the ancestral limb from which the walking branch, or endopod, of the biramous limb evolved, based on morphology (e.g. Boxshall (2004) and references therein) and molecular data (Panganiban et al., 1995, 1997), but the exact mechanism of transition between the two is widely debated. Alternate theories suggest that the exopod flap derived by outgrowth from the lobopod-like limb (Snodgrass, 1938) or from the body wall (Fryer, 1992); or that the biramous limb was the result of a reduction of a polyramous limb (Kukalova-Peck, 1992); or the fusion of two body segments, each of which originally carried a uniramous limb (Em- erson and Schram, 1990). Based on observations from Opabinia suggesting it possessed ventral lobopods, Budd (1996) hypothesised that the arthropod biramous limb evolved through the fusion of a lobopod walking limb and a lateral lobe bearing dorsal gill blades, as is present in this taxon, the “gilled lobopods” from Sirius Passet and the anomalocaridids. Alternately, a different interpretation of Opabinia’s morphology led Zhang and Briggs (2007) to advocate for the “phyllopod scenario” (Olesen et al., 2001), whereby the endopod arose from the margin of Opabinia’s phyllopodous lateral lobe (which, in their view, possesses posterior gill blades) via an intermediate stage possessing a cleft basal region, as seen in Branchiocaris (Briggs, 1976). The new data from Opabinia and the anomalocaridid Hurdia presented in this thesis challenge the assertions made by Zhang and Briggs (2007) and supports the bi-lobed model of biramous limb evolution of Budd (1996).

13 The anomalocaridids

Owing to their large size and inferred predatory habits, the anomalocaridids have received a certain notoriety amongst Cambrian organisms. Their char- acterization is the culmination of over 100 years of research, which began with the description of a single appendage and passed through a convoluted history that eventually led to the reconstruction of one of the most unusual animals in the Cambrian seas. Their body is segmented and bears a series of lateral flaps with associated gills behind a cephalic region consisting of a pair of stalked eyes, a circular mouth apparatus, a pair of frontal appendages and a head shield. The anomalocaridids have been described from most of the major Cambrian lagerstätte, and as such widespread predatory organisms, they undoubtedly played a major role in structuring of the first complex marine animal communities.

History of description from the Burgess Shale The history of description of the anomalocaridids has been complicated by the tendency of their bodies to disarticulate, such that appendages, mouth parts, gills and cephalic carapaces are found as isolated elements much more often than as articulated specimens. This animal group was originally described from Burgess Shale material. The earliest anomalocaridid body parts described are the frontal appendages of Anomalocaris canadensis (Whiteaves, 1892), which were interpreted at the time to be the abdomen and tail of a headless crustacean body (Fig. 3A). The frontal appendage of another anomalocaridid taxon was initially described as “Appendage F” (Fig. 3C) and thought to be the cephalic appendage of the arthropod Sidneyia (Walcott, 1911a; Simonetta, 1963). Walcott described other isolated anomalocaridid body parts as their own taxa, including mouth parts as the medusoid Peytoia nathorsti (Walcott, 1911b) (Fig. 3B), a single whole body specimen (Fig. 3F) as the holothurian Laggania cambria (Walcott, 1911b), and a head shield as the malacostracan Hurdia victoria (Walcott, 1912), (Fig. 3D). Another part of the head shield, Proboscicaris, was described by Rolfe (1962) as an unknown arthropod carapace (Fig. 3E).

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Figure 3. Anomalocaridid body elements. A, Anomalocaris frontal appendage from Mount Stephen trilobite beds. B, Mouthparts, initially described as the medusoid Peytoia (Holotype USNM 57538). C, “Appendage F” frontal appendages (Holotype USNM 57490). D, H-element of Hurdia victoria (Holotype UNSM 57718). E, P- element of Hurdia, originally described as Proboscicaris agnosta (Holotype USNM 139871). F. Whole-body specimen of Laggania cambria with mouthparts (arrow), previously described as a holothurian, polychaete or sponge superimposed with the medusoid Peytoia (Holotype USNM 57555). G. The first whole body specimen of Anomalocaris canadensis described (GSC 75535). Scale bars equal 10 mm.

15 These isolated body parts underwent a series of re-interpretations before being identified as anomalocaridid in origin. The Anomalocaris canadensis appendage was attached to the body of the enigmatic arthropod Tuzoia (Hen- riksen, 1928) although both Simonetta and Delle Cave (1975) and Briggs (1979) suggested instead that it was the appendage of a large, unknown arthropod. “Appendage F” was eventually removed from Sidneyia (Bruton, 1981) but not placed with any other animal. Laggania cambria was rede- scribed as a polychaete worm (Madsen, 1957), or a sponge with the jellyfish Peytoia superimposed on top of it (Conway Morris, 1978). In the latter publication, Peytoia nathorsti was designated as the senior synonym of Lag- gania cambria. The critical work by Whittington and Briggs (1985) recognized the true identity of most of these anomalocaridid parts. Preparation of specimens at GSC (Fig. 3G) and USNM revealed that Anomalocaris and “Appendage F” were actually the frontal appendages of two large animals that had Peytoia for mouthparts. They designated two species, Anomalocaris canadensis (Fig. 4A), which had the original Anomalocaris frontal appendage but no clear mouthparts, and Anomalocaris nathorsti (Fig. 4B), which had “Appendage F” and a clear Peytoia mouth. The latter taxon also included the specimen of Laggania cambria described above (Fig. 3F), which represents a whole body specimen with lateral lobes. In reconstructions of these taxa, Whittington and Briggs (1985) positioned gill-like structures in an internal chamber of the body, an idea opposed by Bergström (1986, 1987), who suggested an alternate reconstruction where lanceolate blades or scales were located on the surface of the lateral lobes. Bergström (1986, 1987) also interpreted mineralized “transverse rods” as thin, segmented ventral appendages. New Burgess Shale material was collected during Royal Ontario Museum (ROM) expeditions, led by D. Collins, during 18 field seasons between the years 1975 and 2001. These expeditions yielded several complete and well- preserved anomalocaridid specimens, allowing a re-examination of the two Burgess Shale anomalocaridid taxa. Collins (1996) found disarticulated assemblages of Anomalocaris canadensis with intact mouthparts identical to the type of Peytoia nathorsti, meaning that Peytoia nathorsti was not a senior synonym of Laggania cambria, but a junior synonym of A. canadensis. The specimens referred to as Anomalocaris nathorsti by Whittington and Briggs (1985) must instead be called Laggania cambria. Laggania is distin- guished from Anomalocaris in that it lacks a tail fan but has a wide head shield, lateral (as opposed to dorsal) eyes, a square mouth part, frontal appendages with long ventral spines, and a rounded body trunk, differences pronounced enough that Collins (1996) felt warranted in maintaining them as separate genera. The same conclusion had been reached by Chen et al. (1994), who had previously suggested renaming Anomalocaris nathorsti to Peytoia nathorsti. Collins (1996) created the order Radiodonta and class Dinocarida to accommodate the anomalocaridids.

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Figure 4. Anomalocaridid diversity in the Burgess Shale. A, Anomalocaris canadensis and B, Laggania cambria reconstructions. Modified from Collins (1996) and drawn by M. Collins. C, Hurdia victoria reconstruction. Modified from Paper I and drawn by M. Collins. D, Reconstruction of ?Laggania frontal appendage. E, Poten- tial Laggania cambria frontal appendage. F, Hurdia victoria and Hurdia triangulata frontal appendage. G, Amplectobelua stephenensis frontal appendage. H, Caryosyn- trips serratus frontal appendage. Scale bars equal 10 mm.

Based on fossil material from the ROM, Collins (1992, 1999) informally described a third Burgess Shale anomalocaridid, Hurdia (Fig. 4C), which is similar in body morphology to the other anomalocaridids but has a promi- nent frontal carapace composed of the triangular Hurdia carapace (now referred to as the H-element) and two Proboscicaris carapaces (P-elements). In fact, Walcott may already have identified a relationship between Hurdia and Proboscicaris almost 100 years ago, because, as Rolfe (1962) points out, Walcott was likely referring to Proboscicaris when he stated that “there are fragments of the carapace of a very large form that possibly may be related to Hurdia” (Walcott, 1912, p. 183). Paper I is a formal description of Hurdia victoria and a discussion of its evolutionary significance, while the details of its morphology and systematics are examined in Paper III. This brings the total of whole-body anomalocaridids known from the Burgess Shale to three, and isolated appendage material from the ROM collection suggests the presence of several more taxa. Amplectobelua, a taxon known from the Chengjiang fauna, and Caryosyntrips, a completely new anomalocaridid, are established in Paper II, along with a more detailed description of a possible frontal appendage of Laggania.

17 The Burgess Shale anomalocaridid Hurdia victoria Two papers in this thesis describe the third Burgess Shale anomalocaridid taxon known from relatively complete full-body preservation. Hurdia victoria is represented by at least 732 specimens, and is the most common anomalocaridid in the Walcott Quarry. Study of this material helps to clarify anomalocaridid systematics and morphology by revealing that previous reconstructions of Anomalocaris and Laggania were partially misled by the inclusion of Hurdia specimens in the studied material of these two genera. As Paper I describes, Hurdia is unique compared to other anomalocaridids in having mouthparts with extra rows of teeth within the central opening (Fig. 5C), two distinct morphs of an “Appendage F”-type frontal appendage (Fig. 4E, F), and a large frontal carapace extending forward from the anterior margin of the body (Fig. 4C, 5A, B, E, F). It consists of a dorsal H-element (formerly the carapace Hurdia) and two lateral P-elements, which are subrectangular and bearing pronounced extensions that join towards the anterior (formerly the carapace Proboscicaris). Gill structures (Fig. 5D) are exquisitely preserved in Hurdia, and consist of a series of lanceolate blades arranged in series, not dissimilar from the gill structures of Opabinia (Bergström, 1986; Budd, 1996). This helps to clarify an enigmatic feature of Laggania, the “transverse rods”, which are highly mineralized linear structures across the centre region of the body trunk, previously interpreted as thin ventral appendages (Bergström, 1986, 1987). The insertion points of the gills have a similar morphology to the transverse rods, making it likely that the latter structures are the points of origin for the gill blades. The affinities of the anomalocaridids has typically been controversial, and various interpretations have regarded them as stem- (Budd, 1993; 2002; Dewel and Dewel, 1998) or crown- (Maas et al., 2004; Chen et al., 2004) group euarthropods, as a sister group to the arthropods sensu lato (Wills et al., 1998; Hou et al., 2006; Hou and Bergström, 2006) or within the cyc- loneuralian worms (Hou et al., 1995). In Paper I, a phylogenetic analysis was performed in order to better understand the evolutionary significance of Hurdia. A branch-and-bound search of 17 taxa placed Hurdia as a sister taxon to a group consisting of Anomalocaris and Laggania, forming a clade in the stem lineage to the euarthropods (Fig. 2). Based on the assumed homology between the frontal carapace of Hurdia and the euarthropod cephalic shield, this phylogenetic position suggests that the origin of head coverings in euarthropods occurred deep in the stem lineage. Hurdia also gives important insight into the origin of arthropod biramous limbs, because it reveals more clearly than before that the morphology of anomalocaridid gills closely resembles that of the setae associated with the outer branches of Cambrian arthropods limbs. This homology supports the theory that the Cambrian biramous limb formed by the fusion of a uniramous walking limb with a lateral lobe structure bearing gill blades (Budd, 1996).

18 Figure 5. Hurdia victoria from the Burgess Shale. A, Dorsolateral specimen (Para- type USNM 274159). B, Camera lucida drawing of USNM 274159. C, Mouthpart with extra rows of teeth (ROM 59260). D, Isolated gill structure (ROM 59261). E, Dorsal specimen (Paratype ROM 59252). F, Lateral specimen (Paratype ROM 49930). Scale bars equal 10 mm. Abbreviations: Ag, anterior gills; Ey, eye; F, frontal appendage; G, gill; H, H-element; M, mouth parts; Re, reticulated structure; P, P- element; S, eye stalk; T, tail lobe; V, mineral vein.

In Paper III, a more detailed description of the morphology of Hurdia allows for clarification of the systematics of this taxon. The diagnostic characteristics of hundreds of disarticulated H- and P-elements, frontal appendages and mouthparts, are analysed with reference to their associations with other body elements and stratigraphic information. Two of the original Hurdia species are confirmed through the use of multivariate morphometric analyses on the outlines of the frontal carapaces. Hurdia victoria Walcott, 1912 has an elongated and narrow H-element, while the Hurdia triangulata Walcott, 1912 H-element is short and wide. Although two morphs of the frontal appendage were described as belonging to Hurdia in Paper I, it was found that Morph B (Fig. 4F) is associated with both Hurdia victoria and Hurdia triangulata. Morph A (Fig. 4E) is tentatively reassigned as the appendage of Laggania cambria. Hurdia dentata Simonetta and Delle Cave, 1975 is considered a nomen dubium.

19 Diversity and global distribution of the anomalocaridids Since the initial description of Anomalocaris and Laggania from the Burgess Shale, several more taxa have been found at this site and other Cambrian localities around the world. In addition to Hurdia (discussed above), two more taxa have been described based on isolated appendage material from the Burgess Shale, the subject of Paper II. Anomalocaridids are also found in China, the USA, Australia and Europe, with a global diversity of eight possible anomalocaridid genera and 14 species. The Chengjiang biota of China, Cambrian Series 2 in age, has produced several taxa of complete anomalocaridids. Anomalocaris saron differs from Anomalocaris canadensis in having a frontal appendage with longer ventral spines and extra lateral spines, and a tail with two long slender furca (Chen et al., 1994). Amplectobelua symbrachiata is an anomalocaridid with a similar body architecture to Anomalocaris, but its frontal appendage is distin- guished by the presence of an exceptionally long ventral spine (Hou et al., 1995). The first occurrence of this taxon outside of Chengjiang is described in Paper II, based on isolated appendages of Amplectobelua stephenensis from the Burgess Shale. A controversial taxon with possible anomalocaridid affinities is Parapeytoia yunnanensis, which may possess biramous limbs and has a frontal appendage with fewer podomeres and spines than other anomalocaridid taxa (Hou et al., 1995). Fragmentary material, usually isolated appendages and mouthparts or rare lateral flaps, has been found at other sites in the Canadian Rocky Mountains (Resser, 1929) and Mackenzie Mountains (Butterfield and Nicholas, 1996), and in southern California (Briggs and Mount, 1982), Utah (Briggs and Robison, 1984; Briggs et al., 2008), Nevada (Lieberman, 2003) and Penn- sylvania (Resser, 1929). Anomalocaris pennsylvanica is a taxon known only from appendages the Cambrian Series 2 Kinzers Formation of Pennsylvania (Resser, 1929). The same is true of Anomalocaris briggsi from the Cambrian Series 2 Emu Bay Shale in Australia (Nedin, 1999) and Cassubia infer- cambriensis from the Terreneuvian Zawiszany Formation in Poland (Dzik and Lendzion, 1988; Masiak and Zylinska, 1994). A possible anomalocaridid appendage is described for the first time from the Cambrian Series 2 Sirius Passet fauna of Greenland in Paper IV.

Anomalocaridid diversity at the Burgess Shale In Paper II, isolated appendage material is used to describe three anomalocaridid taxa from the Burgess Shale, highlighting the importance of disarticulated material for understanding anomalocaridid diversity. The specimens are relatively rare in the collection and full descriptions of the whole- body morphology of these animals await the discovery of new complete specimens.

20 Amplectobelua stephenensis (Fig. 4G, 6A, B) is the first occurrence of this Chengjiang genus outside of China, differing from Amplectobelua symbrachiata in the possession of only 12 podomeres instead of 15, two pairs of large endites instead of one, and paired ventral endites on podomeres 2 to 9 only as opposed to on all of them. All specimens of Amplectobelua stephenensis derive from the “S7” locality on Mount Stephen, and they are generally smaller in size than other described Burgess Shale appendages. A new genus and species, Caryosyntrips serratus (Fig. 4H, 6C, D) consists of a straight and tapering appendage with at least 12 podomeres, each of which has one large spine on the inner surface and several tiny spines lining the outer margin. The twelve specimens come from Fossil Ridge and Mount Stephen. A third type of appendage (Fig. 4D, 6E, F), known only from the “S7” locality on Mount Stephen, is placed tentatively within Laggania, the appendages of which are poorly preserved in whole-body specimens. The morphology is similar to that of the Hurdia Morph A appendage, but it has more ventral spines, including an unusually thin and flexible ventral spine, and a distal end with dorsal spines in similar arrangement to that seen in Amplectobelua. In Paper II, these S7 appendages are placed tentatively in Laggania, but in Paper III, they are described as a possible second species of Laggania. This is because the analysis of stratigraphic distribution in Pa- per III suggests that the Hurdia Morph A appendages are actually the frontal appendage of Laggania cambria, meaning the S7 appendages described in Paper II would belong to a different Laggania species. The frontal appendages of different anomalocaridid taxa from the Burgess Shale are variable in size and shape, which is thought to reflect the different feeding strategies employed by these animals in order to partition prey re- sources and reduce interspecific competition. The “S7” locality has yielded specimens of all five genera found at the Burgess Shale, giving it the highest anomalocaridid diversity of any locality in the world. Diversity of anomalocaridids seems to decrease with time, since the “S7” locality is stratigraphi- cally lower than the Fossil Ridge localities, which gradually decrease in the number of genera present, from four at the lowest sites to two at the upper sites.

21

Figure 6. Diversity of anomalocaridid frontal appendages from the Burgess Shale. A, Amplectobelua stephenensis (Holotype ROM 59495). B, Camera lucida drawing of ROM 59495. C, Caryosyntrips serratus (Holotype ROM 57161). D, Pair of Ca- ryosyntrips serratus frontal appendages (ROM 59501). E, ?Laggania frontal appendage (ROM 59504). F, Camera lucida drawing of ROM 59504.

A possible anomalocaridid from Sirius Passet, Greenland In Paper IV, an appendage with possible anomalocaridid affinities is described from the Sirius Passet biota of North Greenland. Tamisiocaris borealis is known from a single specimen (Fig. 7). The appendage is slightly curved and has a minimum of 17 pairs of straight ventral spines set in a V- shaped orientation. It is broadly similar to Anomalocaris and Cassubia, however no auxiliary spines are visible on the ventral spines and the appendage is unsegmented. The lack of segmentation could be taphonomic, but the surface of the appendage is covered in a fine fabric and the arthrodial membranes and dorsal indentations that delineate appendage boundaries are completely absent. It appears that this appendage was unsclerotized and soft, and

22 perhaps had a similar composition to the Sirius Passet lobopods Keryg- machela and Pambdelurion. Despite differences between the appendages of Tamisiocaris borealis and those of the anomalocaridids, the taxon has been placed questionably within Radiodonta. Similarities in composition between Tamisiocaris and the lobopods could also suggest that it was outside the anomalocaridid clade, oc- cupying an intermediate position in the stem group of the arthropods between the more basal lobopodians and the anomalocaridids. More detailed interpretations of its evolutionary significance await the discovery of complete whole-body specimens.

Figure 7. Tamisiocaris borealis from Sirius Passet, Greenland. A, Holotype MGUH 29154. B, Camera lucida drawing of MGUH 29154. Scale bars equal 10 mm.

23 The lobes and lobopods of Opabinia

As one of the most celebrated and unusual fossils of the Burgess Shale, Op- abinia regalis has been the focus of much debate regarding its morphology and interpretation. It has a segmented body with lateral lobes and gills, and a cephalon bearing five eyes, a ventral mouth, and an elongated proboscis with sharp spines at the terminus. Its affinities were initially uncertain (Whitting- ton, 1975), however Bergström (1986, 1987) suggested it was a relative of the anomalocaridids, a view that has found much support in subsequent analyses (Chen et al., 1994; Budd, 1996; Dewel and Dewel, 1998; Chen and Zhou, 1997; Cotton and Braddy, 2004; Kühl et al., 2009; Paper I). Two main controversies concerning the morphology of Opabinia exist. One problem is determining the exact relationship between the lateral lobes and the gill structures, which has implications for the evolution of the Cam- brian biramous limb. The second issue is whether or not Opabinia possesses lobopod walking limbs, as was suggested by Budd (1996). In Paper V, both of these issues are discussed using previously undescribed fossil material from the ROM and new photographic techniques applied to already described specimens. The lateral lobes of Opabinia’s body were first reconstructed as bearing imbricated sheets of folded, gill-like structures upon their dorsal surfaces (Whittington, 1975) (Fig. 8A), however Bergström (1986) was critical of this model, suggesting instead that the gill-like structures took the form of a series of parallel, lanceolate blades underlying the lateral lobes (Fig. 8B). Budd (1996) elaborated upon this reconstruction, modifying it such that the gill- like structures were attached to the dorsal surface of the lobes (Fig. 8C). Further evidence for the lanceolate blade model of gill reconstruction can be found in the exquisitely preserved gills of the closely related anomalocaridid Hurdia, as described in Papers I and III. A re-examination of Opabinia by Zhang and Briggs (2007) led these authors to suggest that the gill-like structures were posterior fringes of the lateral lobes, instead of being borne atop or below them (Fig. 8D). In Paper V, this model is rejected. Several specimens have lateral lobes with complete, smooth posterior margins (Figs. 9E– G), making it impossible for these to be divided into a posterior fringe. Also, a re-examination of the configuration of the gills and lateral lobes shows clearly that these two structures are always on slightly different stratigraphic levels, such that the former being a continuation of the latter is unlikely. The boundaries between the anterior ‘margins’ of the gills and the lateral lobe to

24 which it is supposedly attached are often uneven and sharply angled, giving the impression of being a broken margin as opposed to the natural attach- ment site of the gills to the lobes. The structure of the gills themselves also does not support the Zhang and Briggs (2007) model, since several specimens show them as distinct units attached along a bar-like structure and/or splayed out, not to mention that similar structures in Hurdia are often found in isolation, as described in Papers I and III. Thus, in Paper V, it is con- cluded that the gills are a series of lanceolate blades attached along to the middle-anterior region of dorsal surface of the lateral lobes.

Figure 8. Reconstructions of lateral lobe and gill structures in Opabinia regalis. A, Highly folded sheet structure on the lateral lobe (Whittington, 1975). B, Lanceolate blades interlayered with the lateral lobes (Bergström, 1986). C, Lanceolate blades on dorsal surface of lateral lobe and attached along anterior region (Budd, 1996). D, Gills as a posterior fringe of the lateral lobe (Zhang and Briggs, 2007). Illustrations redrawn from original publications.

A highly controversial aspect of Budd’s (1996) Opabinia reconstruction was the presence of paired lobopod-like walking limbs extending from the ventral surface of the animal. Several specimens of Opabinia show a series of highly-reflective axial triangles along the length of the body trunk, which had previously been interpreted as musculature (Hutchinson, 1930) or as extensions of the gut or circulatory system (Whittington, 1975), but which Budd (1996) suggests instead to represent “mineralization associated with the internal cavities of lobopod limbs” (Budd, 1996, p. 7). Zhang and Briggs (2007) tried to clarify the identity of these structures using elemental map- ping analysis, concluding that they are undifferentiated gut diverticula extending into the lateral lobes, based on a similar elemental composition shared by the triangular areas and the axial trace of the body. Evidence refut- ing the conclusion of Zhang and Briggs (2007) and supporting the presence of walking limbs is presented in Paper V. The patterns of elemental map- ping are re-examined to show that they are associated with any once fluid- filled body cavity, instead of just those associated with the gut. The photographic techniques employed illustrate that the axial band in Opabinia can be divided into two distinct regions (Figs. 9A, B), a narrow dark band representing the alimentary canal, and a wider band interpreted to be the internal body cavity, with the triangular structures being an extension of the latter (as opposed to the former). We document specimens with a series of paired,

25 elliptical structures that are mineralized or preserved in high relief and associated with the alimentary canal (Figs. 9C, D). These structures are much more likely to represent gut diverticula based on comparisons to other Cam- brian stem-group arthropods such as Kerygmachela (Budd, 1993), Pambde- lurion (Budd, 1998) and Leanchoilia (Butterfield, 2002). Another specimen preserves a fragment of an annulated structure encompassing one of the axial triangles (Figs. 9H, I), perhaps providing physical evidence that the triangular structures represent lobopod limbs. Despite differences in the interpretation of Opabinia’s morphology, phylogenetic analyses consistently place it in an intermediate position between the basal lobopodians and onychophorans, and the anomalocaridids further uptree (compare Zhang and Briggs (2007) with the phylogenetic analysis in Paper I), either in the stem lineage leading to the euarthropods (Budd, 1996; Zhang and Briggs, 2007; Paper I) or in a clade in a sister-group position relative to the arthropods (Wills et al., 1998; Hou and Bergström, 2006). Different interpretations of morphological features can have important implications for evolution and the order of character acquisition, in particular with reference to the arthropods. The reconstruction of the lateral lobes and gills of Opabinia is relevant to the discussion of the evolution of the biramous arthropod limb. The Zhang and Briggs (2007) reconstruction of Op- abinia’s gill structures attaching to the posterior margin of the lateral lobes supports the theory that a segmented endopod arose from the basal region of the lateral lobe (Olesen et al., 2001) and evolved into a jointed, stenopodous limb branch. The implications of this model are that the walking limbs of onychophorans are not homologous to the walking limbs of arthropods. The model suggested by Budd (1996) and elaborated upon in Paper V suggests that the lateral lobes and gills of stem taxa such as Opabinia and the anomalocaridids fuse with a ventral set of lobopod-like limbs to give rise to the Cambrian biramous limb. Thus, the gill structures of Opabinia are homologous to the outer branches of the Cambrian biramous limb, based on their similar morphology as discussed in Paper I, and the ventral limbs were secondarily lost in the anomalocaridids. The homology between onychophoran walking limbs and arthropod walking limbs remains intact, as is also suggested by recent developmental data (Wolff and Scholtz, 2008; Janssen et al., submitted).

26

Figure 9. Opabinia regalis. A, Lateral specimen with alimentary canal (thin black axial line) and gut glands (paired circular structures) contained within body cavity with ventral triangular structures (GSC 40251). White square is close-up in B. B, Axial structures in GSC 40251. C, ROM 59874 with three-dimensionally preserved gut glands D, Gut glands in ROM 59874, coated with ammonium chloride. E-G, USNM 274168, USNM 241479k and USNM 205058 with complete posterior margins of lateral lobes (arrows). H, Central posterior region of YPM 5809 with possible ventral appendage (arrow). I, Camera lucida drawing of YPM 5809. Scale bars equal 5 mm. Anterior to left in all figures except F and G, where anterior is to right. Abbreviations: A, possible appendage; G, gills; L, lateral lobe; R, central region.

27 Conclusions and future perspectives

This dissertation has provided new insight into the morphology, systematics and evolutionary significance of two of the most enigmatic Cambrian taxa, the Anomalocarididae and Opabinia. In both cases, critical information was provided by newly collected specimens from the Burgess Shale and Sirius Passet lagerstätten, emphasizing the importance of continued collecting activities at these and other Cambrian localities. New data was also compiled from previously described specimens, stressing that old material, especially that of enigmatic nature, must also be continually re-evaluated in light of the developments in new methodologies for studying fossil material, and chang- ing opinions on evolutionary patterns. Advanced photographic techniques (Bengtson, 2000) and the application of morphometric statistical methods provided valuable information for clarifying certain troublesome aspects of anomalocaridid morphology and systematics. In this respect, future work on the anomalocaridids could include a re- examination of the first two described anomalocaridid taxa, Anomalocaris and Laggania. In particular, the morphological differences between the mouthparts of different anomalocaridid taxa could be examined quantita- tively with morphometric analysis, and preliminary results (for example, Fig. S2 of Paper I) using advanced photographic techniques on Laggania specimens suggest this approach could help clarify unclear morphological features of the trunk and the relationship between gills and lateral lobes. Other critical taxa with possible anomalocaridid affinities from the Chengjiang fauna of China should be further studied in order to clarify their relationship to the anomalocaridids and their placement in the arthropod stem lineage. For example, full-body specimens of Amplectobelua symbrachiata have yet to be formally described, and the controversial Parapeytoia yunnanensis requires restudy regarding key features such as the form of its mouthparts and lateral lobes. With the above information, a synthesis of anomalocaridid systematics could be attempted incorporating specimens from both the Bur- gess Shale and Chengjiang into phylogenetic analyses to gain a better reconstruction of the arthropod stem lineage.

28 Svensk sammanfattning

Utvecklingen av djuren sträcker sig mer än en halv miljard år tillbaka, till en tid då en betydande evolutionär spridning, den “kambriska explosionen”, gav upphov till nästan alla phyla inom djurvärlden vi känner till idag. Denna spridning är enastående dokumenterad i en mängd fossila lagerstätten, där exceptionell bevaring av djur bestående av mjukdelar tillhandahåller en glimt av komplexa marina ekosystem under den tidigaste utvecklingen av den moderna faunan. Många kambriska fossil förekommer i enigmatiska former vilka väcker livlig debatt angående deras morfologi, släktskap och evolutionära betydelse. En sådan grupp av djur är anomalocariderna, stora förmodade predatorer med en komplex beskrivningshistoria. Den följande avhandlingen presenterar forskning om morfologin, systematiken och den evolutionära betydelsen av anomalocariderna och den närbesläktade Opabi- nia regalis, en annan kambrisk besynnerlighet. De flesta fossil i denna studie kommer från Burgess Shale i Kanada, dis- kuterbart den mest berömda av alla kambriska lagerstätten, och Siriuspasset på Nordgrönland (Fig 1). Anomalocariderna beskrevs först från Burgess Shale för över 100 år sedan. Deras kroppar tenderade att disartikulera, vilket har lett till att extremiteter, mundelar, gälar och huvudsköldar först beskrevs som enskilda delar tillhörande olika djur (se Collin, 1996). Whittington och Briggs (1985) var de första som lade delarna tillsammans och beskrev hela kroppen hos Anomalocaris. Dessa djur har en segmenterad kropp med en serie av breda sidolober med gälar och ett huvud med en cirkulär käft, ett par stora frontala utskott, två själkförsedda ögon, och en huvudsköld (Fig. 4A). Två taxa, Anomalocaris och Laggania (Fig. 4B), som ursprungligen beskrevs av Whittington och Briggs (1985) har studerats igen, och ett flertal nya taxa av Burgess Shale- anomalocarider är här beskrivna (Figs. 4C–H).

Hurdia victoria från Burgess Shale De två första artiklarna i denna avhandling beskriver den tredje Burgess Shale-anomalocariden känd genom bevaring av hela kroppen, Hurdia victoria (Fig. 4C, 5). Undersökning av detta material bidrar till att tydliggöra anomalocaridernas släktskap och morfologi, samt ökar förståelsen för den evolutionära betydelsen av denna grupp.

29 Affiniteten av anomalocariderna har varit utpräglat kontroversiell, och ef- ter ett flertal tolkningar har de ansetts vara stam- (primitiva) eller kron- (hög- re) grupper av euarthropoder, som systergrupp till arthropoderna i generell mening, eller tillhörande Cycloneuralia. Här har en fylogentisk analys ge- nomförts och resultatet placerade Hurdia som systertaxa till en grupp bestå- ende av Anomalocaris och Laggania, vilka skapar en gren på stamlinjen till euarthropoderna (Fig. 2). Hurdia är unik i jämförelse med andra anomalocarider i och med att den har en stor frontal huvudköld som sträcker sig framåt från den främre kanten av kroppen. Huvudskölden var tom och funktionen är inte känd. Hurdia har även exceptionellt bevarade gälstrukturer (Fig. 5D), vilka liknar respiratoris- ka strukturer associerade med benen hos andra kambriska arthropoder. Den- na likhet stöder teorin om utvecklingen av birama extremiteter som föreslår att de formades genom sammanväxning av en uniram gångextremitet och en lateral gälbärande lobstruktur (Budd, 1996). Geometriska morfometriska statistiska analyser tillämpades för att särskil- ja de olika Hurdia-arterna åt. Hurdia victoria har en huvudsköld som är längre och smalare än Hurdia triangulata.

Diversitet och spridning av anomalocarider Åtskilliga nya anomalocarid-taxa är beskrivna genom isolerade frontala utskott från Burgess Shale. Amplectobelua stephenensis (Figs. 4G, 6A, B) är den första förekommande av detta Chengjiang-släkte utanför Kina. Caryo- syntrips serratus (Figs. 4H, 6C, D) är ett nytt släkte och art bestående av ett långt rakt utskott. En tredje typ av utskott (Fig. 4D, 6E, F) känd endast från ”S7”-lokalen på Mount Stephen, är ovisst placerad inom Laggania, vars utskott knappt bevaras hos helkroppsexemplar. Skillnader i storlek och form hos dessa utskott reflekterar troligen olika födostrategier hos dessa djur i syfte att minska konkurrensen arterna emellan. Burgess Shale har den högsta diversiteten av anomalocarider i hela världen. En möjlig anomalocarid har även beskrivits genom ett enda frontalt utskott från Siriuspasset på Nordgrönland. Tamisiocaris borealis är ett böjt utskott med minst 17 par raka ventrala taggar i en V-formad placering (Fig. 6G, H). Den har tydliga likheter med Anomalocaris, dock har den inga taggar på de ventrala taggarna och själva utskottet är osegmenterat. Bristen på segmentering kan inte anses vara tafonomisk då utskottet ligger i en matrix av finkorningt sediment optimalt för bevaring av fina strukturer, men inga bevis för segmentgränser kan ses överhuvudtaget. Utskottet var osklerotise- rat och mjukt, eventuellt hade det en liknande uppbyggnad som Siriuspasset- lobopoderna Kerygmachela och Pambdelurion. Tamisiocaris borealis har osäkert placerats inom Radiodonta, men på grund av den nyss nämnda likhe-

30 ten skulle den även kunna uppta en position i stamgruppen till arthropoderna mellan de mer basala lobopoderna och anomalocariderna.

Morfologin hos Opabinia regalis Opabinia regalis är ett ovanligt Burgess Shale-fossil som hamnat i fokus av mycket debatt angående dess morfologi och tolkning. Den har en segmenterad kropp med sidolober och gälar, och ett huvud med fem ögon, en ventral mun och ett långsträckt utskott med vassa taggar längst ut (Whittington, 1975). Opabinia är närbesläktad med anomalocariderna, men det debatteras om morfologin hos sidoloberna och de associerade gälarna (Fig. 8), samt förekomsten av lobopodlika gångextremiteter (Budd, 1996). En färsk studie av Zhang och Briggs (2007) presenterar en rekonstruktion av Opabinia där de ventrala gångextremiteterna tolkas som tarmkörtlar och gälar anslutna till den bakre kanten av sidoloberna. I denna avhandling visar med hjälp av nya fotograferingstekniker att de ventrala extremiteterna är utväxter av kropps- kaviteten, inte tarmen (Figs. 9A, B), och i ett exemplar är en ringmönstrad struktur bevarade - ett fysiskt bevis för att strukturerna faktiskt är gångben (Figs. 9G-I). Åtskilliga nya exemplar har en serie av parade, elliptiska strukturer associerade med tarmkanalen, dessa representerar mycket mer troligt tarmkörtlar (Figs. 9C, D). Gälarna hos Opabinia visas vara kopplade till den främre regionen av den dorsala sidan av sidoloberna, en rekonstruktion som har viktiga implikationer för utvecklingen av birama ben.

31 Acknowledgements

I was fortunate to have the generous help of many great people throughout the course of my studies. Their support has made this work possible. First and foremost, I thank my main supervisor Graham Budd, for being an unwavering source of support, guidance and encouragement, and for con- stantly challenging me to develop intellectually. I truly appreciate all you have done for me. My co-supervisor Jean-Bernard Caron is also gratefully acknowledged for his contribution to the development of this project and my education, as well as for providing me with the opportunity to work at the spectacular Burgess Shale locality on more than one occasion. I also thank Lars Holmer, my additional supervisor, for his constant support during all stages of this project. I have had the great fortune of working with many wonderful colleagues at Uppsala University. It has been a pleasure spending time on a day-to-day basis with everyone in Palaeobiology at the Department of Earth Sciences, and I thank you all for your encouragement during the past four years. I am especially grateful to John Peel for excellent advice, collaborations and cor- rections to manuscripts. Michael Streng commented on this chapter and provided help over the years with more things that I could possibly name, and Ralf Janssen was great company during numerous Zoonet meetings and trips. Linda Lagebro is thanked for translating my Swedish summary, and more importantly for being a dear friend. I also thank Åsa Frisk and Sebas- tian Willman for answering my recent questions about the defense process. The Evolutionary Organismal Biology research group has always provided a great atmosphere for discussions, which I thoroughly enjoyed over the years, and I thank everyone there for tolerating the occasional presence of an invertebrate paleontologist. Martin Brazeau provided helpful comments on this chapter. Gary Wife and Stefan Gunnarsson are thanked for assistance with the SEM, and Göran Arnqvist is acknowledged for help with statistics. During several lengthy visits to the Royal Ontario Museum, I was warmly welcomed by everyone in the Palaeontology department and I am grateful for the friendly atmosphere and conversations I encountered there. I thank Peter Fenton, Brian Iwama, Ian Morrison, Dave Rudkin and Janet Wadding- ton for providing technical support, as well as Lorna O’Brien for her friendship and for putting up with my frantic last minute email requests to check details in specimens. Des Collins is gratefully acknowledged for collecting, identify, and preparing the vast majority of ROM Burgess Shale specimens.

32 I was fortunate enough to collaborate with several international research- ers, who have each contributed to my academic development in their own ways. Open discussions with Jan Bergström led me to critically examine and develop my own ideas, for which I am grateful. Greg Edgecombe has patiently shared with me a wealth of knowledge on all things arthropod, and I am especially thankful for his contribution to the Hurdia project. I also enjoyed helpful discussions and collaborations with Whitey Hagadorn, Des Collins, and Bob Gaines. Doug Erwin, Jann Thompson and Mark Florence at the Smithsonian National Museum of Natural History, Frederick Collier at the Harvard Museum of Comparative Zoology, and Susan Butts from the Peabody Museum of Natural History provided access to Burgess Shale specimens, as did Jean Dougherty at the Geological Survey of Canada, who is also thanked for her kind hospitality while I was in Ottawa. This research was largely funded by the Marie Curie Research and Train- ing Network ZOONET, through which I had the opportunity to attend Evo- Devo courses and meetings. I am grateful to all members of ZOONET for interesting discussions, and in particular to the other Fellows, for their support, friendship and entertaining times. Additional funding was provided by Uppsala University, and VR research grants awarded to Graham Budd. Tra- vel grants from The Palaeontological Association, The Paleontological So- ciety, The Paleontology Division of the Geological Association of Canada, and the C.F. Liljewalchs Stipend (Uppsala University) allowed me to attend academic meetings. Fieldwork was funded by the Royal Ontario Museum and NSERC grants awarded to Jean-Bernard Caron. Early in my education, I received support and encouragement that undoubtedly influenced my career path in very positive ways. Guy Narbonne nurtured my budding interest in Paleontology during my undergraduate degree at Queen’s University by sharing his enthusiasm with me, and I am very grateful for his support. Jisuo Jin and Alfred Lenz at the University of West- ern Ontario patiently introduced me to Paleontological research during my Masters degree, and I thank them for their encouragement and guidance. I also extend thanks to all my wonderful friends in Sweden who helped make my experiences here so enjoyable: Anna, Ina, Linda and Ralf (and Moltas), The Braz, Tamara, Magnus, Mattias, Claudia, the innebandy crew, ‘the Canadians’, and the palaeogirls. Michi, you have been an endless source of support for me both at work and outside of it, and I truly appreciate you being there for me. To my close friends in Canada, thank you for tolerating my long absences and for making my visits home so entertaining. Finally, I thank my family. Mom, Dad and Sarah - your support has been phenomenal. You’re the best. Thank you.

33 References

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