FIGURE 4-1 - uploaded by James W Hagadorn
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, Arenosicaris inflata from the Furongian Elk Mound Group of Wisconsin-PRI-1013: 1a-c (a at top, c at bottom), three phyllocarids; two partial, and one complete; 2, Ceratiocaris acuminata, PE-664, inset image (YPM 212400) shows a telson with a row of small pits where lateral spines inserted (arrowed); 3, C. bohemica, CNMP L40580, inset image shows a nearly complete carapace valve, CNMP L37712; 4, Warneticaris cornwallisensis, un-numbered telson in the collections of the Czech National Museum, Prague; 5, Warneticaris grata, CNMP L37712, a rare carapace valve showing ornamentation. All scale bars 5 1 cm.
Source publication
A phylogenetic analysis and review of Paleozoic phyllocarid systematics is presented using morphology-based characters from Cambrian to modern taxa. The resulting cladograms of the Phyllocaridasuggest that the suborder Ceratiocaridina as traditionally defined (families Ceratiocarididae and Caryocarididae) isparaphyletic. Caryocarididae is subsequen...
Citations
... In this study we explore the hydrodynamic performance (drag and lift) of caryocaridid carapaces, contrasting them with archaeostracan relatives including the late Cambrian benthic Arenosicaris [24] and Ordovician ceratiocaridids, which are generally considered pelagic [25] or nektobenthic [26]. Thus we determine the role of carapace shape in facilitating a pelagic mode of life in Ordovician caryocaridids. ...
... Arenosicaris carapaces range from 28.6-36.2 mm in length (mean 32.6 mm) [24], the carapace of the first figured specimen of Ceratiocaris angusta is 25 mm in length [40], while those of Ce. silicula are smaller, c. 10-15 mm [25]. Caryocaridids are generally between 10-30 mm in length [21]. ...
... The hydrodynamic performance of the carapaces of the two sampled Ceratiocaris species, C. angusta and C. silicula are broadly comparable to those of most caryocaridids (Figs 3 and 6). Notably a pelagic lifestyle for C. silicula has been previously suggested on account of its narrow and elongate carapace [25], while C. angusta was initially described as a caryocaridid partly on account of its carapace proportions [40]. Collette and Hagadorn [25] posited that the superficial similarities in carapace shape, which lead to the similar drag and lift coefficients from the simulations, could have resulted from ecological convergence to a pelagic mode of life or from a shared ancestral form. ...
The diversification of macroscopic pelagic arthropods such as caryocaridid archaeostracans was a crucial aspect of the Great Ordovician Biodiversification Event, and the plankton revolution. A pelagic mode of life has been inferred for caryocaridids from their common presence in black graptolitic shales alongside carapace morphologies that appear streamlined. However, the hydrodynamic performance within the group and comparisons with other archaeostracans were lacking. Here we use a computational fluid dynamics approach to quantify the hydrodynamic performance of caryocaridids, and other early Palaeozoic archaeostracans including Arenosicaris inflata and Ordovician ceratiocaridids. We show that streamlining of the carapace was an important factor facilitating a pelagic mode of life in caryocaridids, in reducing the drag coefficient and facilitating a broader range of lift coefficients at different angles of attack. However, comparable hydrodynamic performance is also recovered for some ceratiocaridids. This suggests that alongside carapace streamlining, adaptations to appendages and thinning of the carapace were also important for a pelagic mode of life in Ordovician caryocaridids.
... Paleozoic phyllocarids fall into the highly diverse, well-distributed Archaeostraca (Collette and Hagadorn, 2010;Bicknell et al., 2020;Liu et al., 2022Liu et al., , 2023. These crustaceans played key roles as predators or scavengers, feeding on molluscs, arthropods, and carrion (Bergström et al., 1987;Vannier et al., 1997;Bergmann and Rust, 2014;Liu et al., 2022) using large, re-enforced mandibles (Brett and Walker, 2002;Brett, 2003). ...
Trace fossils can illustrate important palaeobiological interactions within a fossil assemblage that body fossils do not record. A group of these trace fossils that showcase feeding ecology, and evidence of predation, are coprolites. Shelly coprolites are useful for documenting records of durophagous predators or scavengers within a substrate. To expand the record of these traces from the lower Paleozoic, here we present 12 shelly coprolites from the Late Ordovician (Katian) Bohdalec Formation of the Czech Republic. These coprolites contain abundant Onnia superba (Bancroft, 1929) fragments with marked breakages across exoskeletal sections. Rarer evidence for gastropods, bivalves, crinoid debris, and another indeterminate shelly material are also observed within the coprolites. While the producer cannot be irrevocably determined, possible options are explored. We propose that larger, co-occurring trilobites and predatory cephalopods likely made the majority of coprolites. Furthermore, large unbiomineralised arthropods, such as phyllocarids and eurypterids are highlighted as possible producers. Continued examination of these trace fossils will highlight when and where similar interactions between trophic levels had occurred.
... Phyllocarids are divided into two classes, an extinct one, middle-late Cambrian to Permian, Archaeostraca (which can be large, have five limb-bearing abdominal somites, a carapace with hinge line, and a telson produced into elongate structure located medially between two lateral furcal rami), and the other one, Permian to recent Leptostraca (which are small, have six limb-bearing abdominal somites, a carapace without a hinge line, and a telson not produced into medial process between lateral furcal rami). These arthropode fossils occur in intertidal facies, an ecological niche that a substantial portion of modern leptostracan species still recently inhabit [ 15 ]. ...
Palaeozoic phyllocarid crustaceans and pterygotid eurypterids from the Gethicum Zone of Carpatho-Balkanides in Serbia are herein documented for the first time. The described and illustrated phyllocarid remains consisting of a coupled telson and furcal rami were collected from upper Silurian (?) shale. This specimen is fragmented and poorly preserved to allow identification to species level, and has thus been assigned to Ceratiocaris sp. indet. The eurypterids are represented by part of a fragment, showcasing the distinct scale-like ornamentation of the pterygotids, and a segment of a walking leg. The pterygotid remains were found together with a few graptolite taxa (Monograptus sudeticus Porębska, “M.” microdon microdon Reinh. Richter, Neomonograptus aequabilis (Přibyl), Uncinatograptus hercynicus (Perner), U. subhercynicus (Willefert)), which are characteristic for the lower part of the U. hercynicus graptolite zone (Lochkovian, Lower Devonian). There are at least two species in this stratigraphic interval that correspond to the pterygotid eurypterids remains from Serbia – Acutiramis perneri Chlupáč and Pterygotus cf. barrandei Semper, but without the chelicerae, it is impossible to make precise identification. These new finds from eastern Serbia, together with the Bulgarian ones, are a further confirmation of peri-Gondwanan origin of pre-Variscan Palaeozoic sediments in the West Srednogorie Unit in Bulgaria and in the Gethicum Unit in Serbia.
... Phyllocarida Packard, 1879, which have long been considered a monophyletic group, may represent the most basal group of malacostracans (Collette & Hagadorn, 2010a;Rolfe, 1969). Unlike other malacostracans, phyllocarids share the presence of seven abdominal segments or pleomeres (Dahl, 1987). ...
... Hegna et al., 2020;Hirata et al., 2019;Hirata & Kikuchi, 2022;Mees, 2015;Moreira et al., 2021;Schram & Malzahn, 1984;Song et al., 2017) and extinct (i.e. Palaeozoic) Archaeostraca (Collette & Hagadorn, 2010a;Rode & Lieberman, 2002). Phylogenetic analyses suggest that the archaeostracans are probably the sister group of leptostracans (Wolfe et al., 2016), but may also be paraor polyphyletic (Hegna et al., 2020). ...
... Phyllocarid fossils occur as flattened remains in the mudstones and shales. Torsvik & Cocks, 2017; fossil data from Collette & Hagadorn, 2010a;Collette & Rudkin, 2010;Jones et al., 2015a, b;P'an, 1962; and this paper.) B, fossil sites of phyllocarids and C, the phyllocarids-bearing stratigraphical column of Tiejishan section from Wuhan, Hubei. ...
In the Silurian, the suborder Caryocaridina of the phyllocarids was replaced by the suborder Ceratiocaridina as the dominant group. The latter did not achieve a global distribution until the late Silurian. In the early Silurian, ceratiocarids were a little-diversified group and palaeogeographically restricted mainly to Laurussia. Although previous studies have mentioned phyllocarid fossils from the Silurian of the South China plate, these have never been systematically described. This study describes three genera and four species (including one new genus, two new species, and two undetermined species), i.e. Cugocaris future gen. et sp. nov., Gonatocaris wuhanensis sp. nov., Gonatocaris sp. and Warneticaris sp., based on 38 phyllocarid specimens from the Fentou Formation in Wuhan. Cugocaris future gen. et sp. nov. is distinguished by an elliptic carapace ornamented by sinuous and anastomosing longitudinal striae, a styliform telson, and slender furcal rami. Gonatocaris wuhanensis sp. nov. is characterized by anastomosing longitudinal striae on the carapace, greatest carapace width in the centre of the carapace, and three different types of ornamentations on the surface of the abdominal segments. A phylogenetic analysis of 41 Palaeozoic phyllocarids reveals that Gonatocaris, Rolfecaris and Cugocaris gen. nov. form a monophyletic group. Thus, a new family, Gonatocarididae fam. nov., is proposed. This family is characterized by a carapace lacking median dorsal plate, rostral plate, a well-developed anterior carapace horn, and ornamented with prominent raised, wavy, anastomosing longitudinal striae. A palaeoecological analysis suggests that G. wuhanensis was a fairly active swimmer, while C. future was necto-benthic with relatively weak swimming abilities. This study not only extends the known morphological range of phyllocarids, but also has implications for the functional morphology, taxonomy, and evolution of archaeostracans as a whole. http://zoobank.org/urn:lsid:zoobank.org:pub:4B6B60B6-19F1-4A58-AC65-3B1AEE5D0939
... Discussion: Thalassinoides specimens are generally considered dwelling (domichnia) and feeding (fodinichnion) structures of decapod crustaceans. Palaeozoic occurrences of Thalassinoides predate the oldest known body fossils of burrowing decapods and, therefore, other producers have been suggested (Myrow, 1995;Carmona et al., 2004;Buatois & Mángano, 2011): Trilobites, worm-like organisms (Myrow, 1995;Carmona et al., 2004) and phyllocarids which are known since the Cambrian and were common through the Palaeozoic (Collette & Hagadorn, 2010). Thalassinoides occurs in a wide environmental range but mostly in shallow marine settings, related to high availability of oxygen amount (e.g., Knaust, 2017;El-Sabbagh et al., 2017;Vinn et al., 2020). ...
The mixed siliciclastic-carbonate Teferguenite Formation (Pragian-Emsian) occurring in the Ougarta Range (SW
Algeria) is subdivided into two members: the Lower Member consists of three thick limestone beds (namely
bars) intercalated with clay and sandstone deposits; the Upper Member is composed of intercalated sandstone,
silty limestone and clay. Sedimentological data and their associated ichnological content were used for the
interpretation of depositional settings. Lithofacies analysis evidenced that the studied succession was deposited
in a shallow marine (shoreface to upper offshore), storm-influenced setting. Ichnological data document moderate
to high bioturbation intensity, while ichnodiversity consists of 29 ichnospecies, namely: Arenicolites isp., Arthraria
antiquata, Arthrophycus isp., Bergaueria perata, Bolonia lata, Chondrites targionii, Cochlichnus anguineus, cf.
Cruziana isp., Cruziana rusoformis, Diplichnites isp., Gordia marina, Helminthopsis abeli, Hormosiroidea isp.,
Lockeia siliquaria, Megagrapton irregulare, Monomorphichnus cf. multilineatus, Neonereites uniserialis, N.
multiserialis, Nereites isp., Phycodes isp., Protovirgularia isp., Rusophycus isp., Thalassinoides isp., T. suevicus,
Palaeophycus isp., P. striatus, P. tubularis, and Zoophycos ispp. (types A and B). These trace fossils are gathered
in three trace-fossil assemblages: the Palaeophycus-Thalassinoides-Arenicolites ichnoassemblage (PTAI),
Cruziana-Thalassinoides ichnoassemblage (CTI) and Neonereites-Zoophycos ichnoassemblage (NZI). The tracefossil records within the Teferguenite Formation is represented by different behaviours, corresponding to the
archetypal Cruziana ichnofacies occurring from the shoreface to upper offshore zones.
... Thalassinoides specimens are generally considered dwelling (domichnia) and feeding (fodinichnion) structures of decapod crustaceans. Palaeozoic occurrences of Thalassinoides predate the oldest known body fossils of burrowing decapods and, therefore, other producers have been suggested (Myrow, 1995;Carmona et al., 2004;Buatois & Mángano, 2011): Trilobites, worm-like organisms (Myrow, 1995;Carmona et al., 2004) and phyllocarids which are known since the Cambrian and were common through the Palaeozoic (Collette & Hagadorn, 2010). Thalassinoides occurs in a wide environmental range but mostly in shallow marine settings, related to high availability of oxygen amount (e.g., Knaust, 2017;El-Sabbagh et al., 2017;Vinn et al., 2020). ...
The mixed siliciclastic-carbonate Teferguenite Formation (Pragian-Emsian) occurring in the Ougarta Range (SW Algeria) is subdivided into two members: the Lower Member consists of three thick limestone beds (namely bars) intercalated with clay and sandstone deposits; the Upper Member is composed of intercalated sandstone, silty limestone and clay. Sedimentological data and their associated ichnological content were used for the interpretation of depositional settings. Lithofacies analysis evidenced that the studied succession was deposited in a shallow marine (shoreface to upper offshore), storm-influenced setting. Ichnological data document moderate to high bioturbation intensity, while ichnodiversity consists of 29 ichnospecies, namely: Arenicolites isp., Arthraria antiquata, Arthrophycus isp., Bergaueria perata, Bolonia lata, Chondrites targionii, Cochlichnus anguineus, cf. Cruziana isp., Cruziana rusoformis, Diplichnites isp., Gordia marina, Helminthopsis abeli, Hormosiroidea isp., Lockeia siliquaria, Megagrapton irregulare, Monomorphichnus cf. multilineatus, Neonereites uniserialis, N. multiserialis, Nereites isp., Phycodes isp., Protovirgularia isp., Rusophycus isp., Thalassinoides isp., T. suevicus, Palaeophycus isp., P. striatus, P. tubularis, and Zoophycos ispp. (types A and B). These trace fossils are gathered in three trace-fossil assemblages: the Palaeophycus-Thalassinoides-Arenicolites ichnoassemblage (PTAI), Cruziana-Thalassinoides ichnoassemblage (CTI) and Neonereites-Zoophycos ichnoassemblage (NZI). The trace-fossil records within the Teferguenite Formation is represented by different behaviours, corresponding to the archetypal Cruziana ichnofacies occurring from the shoreface to upper offshore zones.
... The position of Cephalocarida and Branchiopoda remains uncertain. All of these Pancrustacea lineages are very old, as evidenced by the fact that there were already malacostracans (Collette and Hagadorn 2010) and branchiopods (Waloszek 1993) present in the Cambrian era. Phylogenetic analysis has allowed sci entists to confirm this (Regier et al. 2005), which implies that Pancrustacea has a truly ancient history with numerous lineages, a large part of which has probably disappeared. ...
... The telson to furca ratio varies greatly within the species [ 5,19 ], but there are calculated averages for the different species [ 1 ]. This ratio for C. bohemica is 1.59, and for C. papilio -1.8 [ 1 ]. ...
... The telson to furca ratio varies greatly within the species [ 5,19 ], but there are calculated averages for the different species [ 1 ]. This ratio for C. bohemica is 1.59, and for C. papilio -1.8 [ 1 ]. The Wenlockian species C. crispus [ 7 ] from Algerian Sahara have a ratio between 2.2 and 2.4. ...
Paleozoic phyllocarid crustaceans are established for the first time in Southeastern Europe. The phyllocarid remains consisting of telsonal part and furcal rami from Wenlockian shale of the Svoge Unit (West Bulgaria) are described and illustrated. This specimen is too poorly preserved to allow identification to the species level, and has thus been assigned to Ceratiocaris sp. These occurrences nevertheless suggest that phyllocarids may be a common component of assemblages present in shale facies since the Silurian in the peri-Gondwanan Europe.
... The position of Cephalocarida and Branchiopoda remains uncertain. All of these Pancrustacea lineages are very old, as evidenced by the fact that there were already malacostracans (Collette and Hagadorn 2010) and branchiopods (Waloszek 1993) present in the Cambrian era. Phylogenetic analysis has allowed sci entists to confirm this (Regier et al. 2005), which implies that Pancrustacea has a truly ancient history with numerous lineages, a large part of which has probably disappeared. ...
... However, but the gash extending from the pygidial margin cannot be ascribed to this form of predation. Options for this are the phyllocarids with large frontal appendages (Fig. 5e) (Collette and Hagadorn 2010) or the pterygotid sea scorpions, such as Acutiramus perneri Chlupáč, 1994; both of which are known to Lochkov Formation (Fig. 5c, d). We suggest that the eurypterids, armed with large raptorial frontal appendages and gnathobases on proximal appendage sections, were the likely groups to have produced the extreme injury across the pygidium. ...
Trilobite malformations are often ascribed to failed predation and represent key evidence for Paleozoic arthropod predator–prey systems. A large number of malformed trilobites are known to Cambrian-aged deposits and have recently been discussed at length. Conversely, most post-Cambrian records are noted as anecdotal points within larger taxonomic works. To expand the consideration of post-Cambrian injured trilobites, we report two malformed Ogygiocarella debuchii specimens from the Middle Ordovician of Wales and a heavily malformed Spiniscutellum umbelliferum specimen from the Early Devonian of the Czech Republic. These specimens are considered to represent records of failed predation. In considering these specimens, we explore possible injury-making groups, in particular noting that S. umbelliferum was likely prey for multiple apex predators. Continued examination of injured trilobites represents the main direction for uncovering how this iconic group of biomineralised arthropods interacted with higher tropic levels within Paleozoic foodwebs.
