Multiple-Rusophycus (Arthropod Ichnofossil) Assemblages and Their Significance

Rusophycus is an ichnogenus comprising shallow burrows generally attributed to trilobites and other arthropods. The paleoethological interpretation of these structures is not conclusively known; workers variously have attributed the ichnogenus to feeding, resting, hiding/escape, hunting or nesting behaviors. Rusophycus morphology varies from unornamented, bilobed forms to forms that preserve details of ventral anatomy of the trace maker. Rusophycus occurs as single or multiple impressions. Some examples of multiple-Rusophycus assemblages are clearly the result of the activity of a single animal (e.g., serially arranged impressions of the same size), but others represent the activity of several individuals. Associated traces representing multiple individuals are especially interesting, for they may give evidence of complex behavior, for example, interactions between the trace makers. Some of the multiple-trace assemblages show alignment (congruent anterior-posterior orientation of the individuals) suggesting rheotactic behavior; other assemblages comprise randomly oriented traces. The difference between the aligned and randomly oriented assemblages may reflect differences in the current energy and amount of available food, and may also suggest the following feeding modes for these benthic-feeding arthropods: (1) alignment with head into the current in resource-poor environments, (2) orientation with head at an oblique angle to the current in high-energy, resource-poor environments, and (3) random orientation in low-energy, resource-rich environments.     

Rusophycus carleyi (James, 1885), Trace Fossils from the Lower Ordovician of Southern Morocco,and the Trilobites that Made Them

Thin-bedded, pyrite-rich, fine sandstones and mudstones of the Floian-Dapingian Upper Fezouata Formation contain abundant trace fossils Rusophycus carleyi in close association with a species of the asaphid trilobite Asaphellus. The sizes and shapes of this trilobite and the traces match closely. Five specimens have even been found where an articulated specimen of Asaphellus appears to be directly located over a specimen of Rusophycus carleyi within a thin bed of sandstone, suggesting that the trilobite animal may have been trapped on top of a trace that it had just made. Such intimate associations between a putative tracemaker and a trace are rare in the fossil record and particularly rare for Trilobita. The number of coxal impressions that form part of R. carleyi, eleven, matches the number expected for an asaphid trilobite (one for each of eight thoracic segments and one for each of three post-oral cephalic appendages). Impressions of the hypostome, thoracic tip impressions, cephalic margin, and pygidial margin in a few of the traces also match those of this asaphid trilobite. R. carleyi has been found in Ordovician strata of other parts of the world in association with asaphid trilobites.     

An opportunistic Upper Ordovician trilobite assemblage from Missouri

The trilobite Ampyxina bellatula (Savage) occurs in monospecific accumulations within dolomitic strata of the shaly Upper Ordovician Maquoketa Group of northeastern Missouri. A dolomitic stratum containing 97 complete individuals and considerable amounts of pyrite exhibited no associated benthic megafauna. The strata appear to be equivalent to the Scales Formation of the Maquoketa Group of northern Illinois and eastern Iowa which have been interpreted as having been deposited in a muddy environment with severe circulation restrictions. The presence of the monospecific trilobite association in what otherwise would be an unfossiliferous stratigraphic sequence is interpreted as being the result of opportunistic strategy by this trilobite in a resource-rich unstable environmental setting. The association was apparently cat-astrophically buried, thus preserving the age structure of the holaspid population. A size-distribution histogram of the holaspid population indicates that reproduction may have occurred in periodic bursts with lesser reproductive events occurring during the intervening periods. Holaspid survivorship information indicates that mortality was low during the early holaspid stage and increased with age. Anomalous single bed trilobite accumulations of various age rocks have been documented by numerous authors. These may in part be analogous to the Ampyxina occurrence in the Maquoketa Group of Missouri, and be attributable to opportunistic strategies.     

Late Ordovician and early Silurian Proetida (Trilobita) from north-western and central Europe

Species belonging to genera of three families of the Proetida (Proetidae, Rorringtoniidae and Toernquistiidae) are described from the upper Ashgill of southern Sweden, and the same or related species noted from the adjacent parts of northern and central Europe. Proetidae are represented by Paraproetus brevifrons (Angelin), Ogmocnemis irregularis Kielan, Ogmocnemis jaanussoni sp. nov. and Parvigena putealis sp. nov.; it is argued that all belong to the Cornuproetinae, not Tropidocoryphinae where they were previously classified. Ogmocnemis is resurrected for some Upper Ordovician species included previously in Decoroproetus . The rorringtoniid genera Rorringtonia and Madygenia are recognised for the first time in southern Sweden, and it is argued that these are probably closely related, and both are included in the same family; both are recorded for the first time from Silurian (Llandovery) strata. Zetaproetus is claimed to be a junior subjective synonym of Rorringtonia. Solariproetus , first described from north-west China, is assigned to the Toernquistiidae as a late, derived member, and S. prosthemesos sp. nov. is described from late Rawtheyan strata in the Holy Cross Mountains, Skåne and the English Lake District. Comments are given on the stratigraphical and palaeogeographical distribution of these taxa.     

An Early Ordovician Trilobite assemblage from the Lashkarak Formation, Damghan area, northern Iran

A low diversity Lower Ordovician trilobite assemblage is described from the middle part of the Lashkarak Formation at Simeh-Kuh, northwest of Damghan, northern Iran. Co-occurrence with conodonts of the lowermost Paroistodusproteus Biozone suggests an uppermost Tremadocian age. This assemblage includes four taxa, including one new genus and two new species, Damghanampyxginteri nov. gen. nov. sp. and Asaphellusfecundus nov. sp. The third diagnosable species from the formation is Taihungshaniamiqueli (Bergeron, 1894) which has previously been reported from southern France and the Turkish Taurides, suggesting north Gondwanan faunal affinities.     

Arthropod Trace Fossils of the Upper Lower Cambrian Gog Group,Southern Rocky Mountains of Canada

The upper lower Cambrian Gog Group in the southern Rocky Mountains of Canada displays a high diversity and abundance of arthropod traces. Four ichnogenera, Cruziana, Diplichnites, Monomorphichnus, Rusophycus, and “Indeterminate Arthropod Scratches” are discussed, with a total of 17 different ichnospecies, as follows: Cruziana billingsi Fillion and Pickerill, 1990, Cruziana irregularis Fenton and Fenton, 1937, Cruziana jenningsi Fenton and Fenton, 1937, Cruziana navicella Fenton and Fenton, 1937, Cruziana omanica Seilacher, 1970, Cruziana pectinata Seilacher, 1994, Cruziana penicillata Gibb, Chatterton, and Pemberton, 2009, Cruziana plicata Crimes, Legg, Marcos, and Arboleya, 1977, Cruziana tenella Linnarsson, 1871, Diplichnites twelvetreesi (Chapman, 1928), Monomorphichnus bilinearis Crimes, 1970b, Monomorphichnus lineatus Crimes, Legg, Marcos, and Arboleya, 1977, Rusophycus eutendorfensis (Linck, 1942 Linck, O. 1942. Die spur Isopodichnus. Senckenbergiana, 25: 232–255. [Google Scholar]), and Rusophycus unilobus (Seilacher, 1970 Seilacher, A. 1970. Cruziana stratigraphy of “non-fossiliferous” Palaeozoic sandstones. In Crimes, T. P. and Harper, J. C. (eds.), Trace Fossils. Seel House Press, Liverpool, 447–476. [Google Scholar]). Three new ichnospecies, i.e., Cruziana caputinclinata isp. nov., Rusophycus subnotous isp. nov., and Diplichnites obliquus isp. nov., are proposed, described, and illustrated. The ichnofauna collected from three localities (Lake O'Hara, Mount Babel, and Redoubt Mountain) are predominantly Cruziana. Evidence of nutrient-rich substrates containing microbially induced sedimentary structures and the ichnospecies present confirm that the trace fossil bearing strata are part of the Cruziana ichnofacies and were deposited in a fully marine basin between fair weather wave base and storm wave base.     

Lepidocoleus cf.ulrichi (Machaeridia) from the Trenton Group (Ordovician) of Ontario (Canada)

A specimen of the machaeridianLepidocoleus cf.ulrichi described herein focuses attention on taxonomical difficulties with extremely dorso-ventrally elongated sclerites found in some lepidocoleid taxa.Lepidocoleus ulrichi Withers, 1926 andLepidocoleus sigmoideus Withers, 1926 exhibit very similar appearances and prove indistinguishable based onWithers 1926 criteria. The key to separating taxa as similar as these may instead lie in the most juvenile stages of growth.     

The trace fossil Sinusichnus from the Upper Ordovician of Anticosti Island,eastern Canada

Branched, sinusoidal burrows assigned to Sinusichnus cf. seilacheri Knaust et al. 2016; occur in Upper Ordovician (Hirnantian) carbonates on Anticosti Island. These are abundant and monospecific at the base of the Oncolite Bed of the uppermost Ellis Bay Formation, marking a regional discontinuity surface during the multiphase Ordovician mass-extinction event. In contrast to Mesozoic and Cenozoic records of Sinusichnus, commonly attributed to the activity of decapod and isopod crustaceans, the burrow systems from Anticosti Island were probably produced by other arthropods of unknown affinity. A combined dwelling, locomotion and feeding behaviour of their trace maker is assumed. This is the oldest record of the ichnogenus Sinusichnus, previously only known since the Triassic, and its first evidence from North America.     

Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

Bryozoans as taphonomic engineers,with examples from the Upper Ordovician (Katian) of Midwestern North America

A combination of encrusting calcitic bryozoans and early seafloor dissolution of aragonitic shells recorded in the Cincinnatian Series of the upper Midwest of North America allowed the preservation of abundant moulds of mollusc fossils bioimmured beneath the attachment surfaces of the bryozoans. We here call this preservational process ‘bryoimmuration’, defined as a bryozoan‐mediated subset of bioimmuration. The bryozoans moulded very fine details of the mollusc shells, usually with more accuracy than inorganic sediment moulds. Most of the bryozoans are heterotrypid trepostomes with robust low‐Mg calcite skeletons. The molluscs are primarily bivalves, gastropods, nautiloids and monoplacophorans with their originally aragonitic shells now dissolved. Many of the encrusting bryozoans are so thin and broad that they give the illusion of calcitic mollusc shells clinging to the moulds. Some molluscs in the Cincinnatian, especially monoplacophorans and epifaunal bivalves, would be poorly known if they had not been bryoimmured. Unlike internal and external moulds in sediment, bryoimmured fossils could be transported and thus record aragonitic faunas in taphonomic assemblages (e.g. storm beds) in which they would otherwise be rare or absent. In addition, bryoimmurations of aragonitic shells often reveal the ecological succession of encrustation on the shells by exposing the earliest encrusters and borings that were later overgrown. Bryoimmuration was common during the Late Ordovician because the calcite sea at the time quickly dissolved aragonitic shells on the seafloor before final burial, and large calcitic bryozoans very commonly used molluscs as substrates. Bryoimmuration is an important taphonomic process for preserving aragonitic faunas, and it reveals critical information about sclerobiont palaeoecology. Several Cincinnatian mollusc holotypes are bryoimmured specimens. Bryozoans involved in bryoimmuration enhance the preservation of aragonitic fauna and thus act as taphonomic engineers.     

Fustiglyphus annulatus from the Ordovician of Ontario,Canada, with a systematic review of the ichnogenera Fustiglyphus Vialov 1971 and Rhabdoglyphus Vassoievich 1951

The ichnogenera Rhabdoglyphus Vassoievich 1951 and Fustiglyphus Vialov 1971, both a source of taxonomic confusion and inconsistent usage, are re‐examined. Many specimens of Fustiglyphus have been incorrectly described as Rhabdoglyphus, both prior and subsequent to the formulation of the former by Vialov in 1971. Updated synonymies for both ichnogenera and their ichno‐species indicate that two ichnospecies of Rhabdoglyphus can be distinguished, namely R. grossheimi Vassoievich 1951 and R. spinosus Ksiazkiewicz 1977, while F. annulatus Vialov 1971 is the only acknowledged ichnospecies of Fustiglyphus. The significant feature distinguishing the two ichnogenera is regarded as the presence of invaginated segments on Rhabdoglyphus as opposed to well‐defined rings or knots with no invagination on Fustiglyphus.

Although Spongolithus annulatus Fritsch 1908 is technically the senior synonym of Fustiglyphus annulatus, it is recommended that the latter name continue to be used in the interests of maintaining nomenclatural stability. Previously unreported specimens of Fustiglyphus annulatus from the Ordovician of Ontario are described and illustrated; one of these suggests that the swellings in Fustiglyphus possibly represent brood chambers of the producing‐organism(s). We also suggest that Fustiglyphus "retreated”; from onshore to deep‐water offshore environments at the end of the Paleozoic.     

A New Endosymbiont in Late Ordovician Tabulate Corals from Anticosti Island,Eastern Canada

Cylindrical tubes of the trace fossil Chaetosalpinx occur within the skeletal walls of Late Ordovician sarcinulid tabulate corals preserved on Anticosti Island. A large host-specific embedment structure, Chaetosalpinx rex isp. nov., is described from the reefal sarcinulid genus Columnopora that occurs in (Rawtheyan, Ashgill) coral-stromatoporoid patch reefs of the upper Vauréal Formation (Mill Bay Member). While locally abundant in Columnopora, this newly described embedment structure is absent in other frame-builders in the reefs.     

Early heliolitine tabulate corals from the Sandbian (Upper Ordovician) in the Yunnan-Sichuan border area,Southwest China

Heliolitine tabulate corals are extremely rare in pre-Katian (Upper Ordovician) rocks worldwide, hindering the understanding of their origin and early evolution. Here, we systematically study new heliolitine material from the upper member (Sandbian) of the Huadan Formation in the Yunnan-Sichuan border area, Southwest China, which includes Protaraea ningnanensis, Plasmoporella kiaeri, Wormsipora sp. and Heliolites spinosus. The findings indicate that Protaraea and Heliolites are ancestral forms of coccoserids and typical tubular heliolitines, respectively, and that the absence of trabecular septa is probably a primitive feature of plasmoporellids. The refined Ordovician stratigraphy of this area implies that the coccoserid Protaraea ningnanensis Lin, 1965 from the lower member (late Darriwilian) of the Huadan Formation is the earliest known heliolitine, which further indicates that all heliolitines might have evolved from Protaraea. The comparison of Protaraea with other early tabulate corals also suggests an independent heliolitine origin from auloporinids and sarcinulinids.     

Silicified Upper Ordovician trilobites from Pai‐Khoi,Arctic Russia

Abstract: A collection of silicified trilobites extracted from samples obtained by F. Nansen from the Upper Ordovician of Khabarovo, Pai Khoi, Russian Arctic is dominated by two new species of the proetide Lasarchopyge, a genus known hitherto only from the Argentine Precordillera. Comparison of Lasarchopyge with Scharyia reveals a range of common characters that support inclusion of both genera in the same family. The rest of the fauna comprises a metagnostid (Trinodus elspethi), an asaphid (Isotelus sp.), two remopleuridids (Remopleurides cf. caelatus and R. sp. nov. A), an aulacopleurid (Harpidella triloba), a lichid and an odontopleurid (Apianurus cf. barbatus), the last two being represented by very fragmentary material. T. elspethi, H. triloba, R. cf. caelatus and A. cf. barbatus are common to or closely resemble taxa from the Edinburg Formation, Virginia. Closely related taxa are present also in the Esbataottine Formation, NW Territories, Canada, and in the Albany Group, Girvan district, Scotland. All of these occurrences are of early Caradoc (Sandbian) age, and most lay at palaeolatitudes of 20°–30° south; all are interpreted as having been deposited on the outer shelf, which is consistent with their distribution.     

Upper Ordovician bryozoans from the Xiazhen Formation of Yushan,northeastern Jiangxi,East China

The bryozoan fauna from the Xiazhen Formation (Katian, Upper Ordovician) of northeast Jiangxi Province, southeast China is reported here. Seventeen species of bryozoans belonging to fifteen genera and four orders are identified: Homotrypa yushanensis, Homotrypa sp., Prasopora yushanensis, Trematopora sp., Monotrypella sp., Rhombotrypa sp., Orbignyella sp., Constellaria jiangxiensis, Constellaria sp., Stictopora nicholsoni, Trigonodictya parvula, Ptilodictya ensiformis, Stictoporella sp., Pseudopachydictya sp., Nematopora sp., Arthrostylidae sp. indet., and Chasmatoporidae sp. indet. Four of these genera have been reported previously but nine genera (Trematopora, Monotrypella, Rhombotrypa, Orbignyella, Trigonodictya, Ptilodictya, Stictoporella, Pseudopachydictya, and Nematopora), one rhabdomesine and one fenestrate are found for the first time in the Late Ordovician strata of South China. Our palaeogeographical analysis suggests that the bryozoan association is typical for the Katian, which is mostly widespread in Laurentia, Siberia, Baltica and Mediterranean, and displays palaeobiogeographical relationships to the Laurentia–Siberia Province.     

An intermittent mode of formation for the trace fossil Cruziana as a serial repetition of Rusophycus: the case of Cruziana tenella (Linnarsson )

Cruziana is one of the most recognizable trace fossils ascribed to arthropods. It ranges throughout the Phanerozoic and encompasses a diverse set of morphologies. The distinct features of Cruziana have incited fierce debate regarding its mode of formation. Here, we discuss critical aspects of trace fossil formation, namely epibenthic versus endobenthic origin and the ethology of the producer. Cruziana has largely been interpreted as a continuous ploughing trace fossil. It has been suggested, however, that at least some Cruziana could be structures resulting from the concatenation of Rusophycus‐type elements, although this claim remains unexplored. Cruziana tenella from the lower Cambrian of south‐central Sweden illustrates this intermittent mode of formation with a series of Rusophycus eutendorfensis leading into vertically undulating Cruziana that, at end stages of development, reflect a relatively equal depth distribution throughout the trace fossil and a great number of intergrading morphologies. In this study, a morphological evaluation of the intergrading morphologies of Cruziana tenella and Rusophycus eutendorfensis and a short morphometric analysis of the elements comprising Cruziana tenella suggests that at least in some cases Cruziana could be formed in intervals, as the serial overlap of distinct shallow Rusophycus could produce an apparently continuous cruzianaeform morphology. A comparison with possible evidence of intermittent formation on Cruziana semiplicata is made to illustrate the possibility of extending this mode of formation to larger Cruziana. An argument for the rise in the early Cambrian of a primitively intermittent mode of formation is made on the basis of energy efficiency.     

The Upper Tremadocian (Ordovician) graptolite Bryograptus: taxonomy, biostratigraphy and biogeography

Abstract:  The taxonomy, biostratigraphical and palaeogeographical distribution of the Lower Ordovician graptolite genus Bryograptus is evaluated. Bryograptus is recognized as a distinct triradiate anisograptid with a multiramous, pendent rhabdosome. The species of the genus Bryograptus can be interpreted as shallow water faunal elements with a strongly limited biogeographical distribution to the Atlantic Faunal Realm. Bryograptus is restricted to a narrow interval in the Upper Tremadocian, the Bryograptus Biozone of Scandinavia and South America (Argentina), making it a taxon with a high potential for precise biostratigraphical correlations. The proximal end development can be used to differentiate the genus Bryograptus from other pendent multiramous graptoloid genera with a homoplastic rhabdosome development. Characteristics of the proximal end development and structure easily differentiate these genera in relief specimens, but not in flattened material.     

新疆柯坪和甘肃平凉上奥陶统底部附近的牙形刺

新疆柯坪大湾沟和甘肃平凉官司庄剖面是Nemagraptusb gracilis带及上奥陶统底界的重要剖面,研究这两剖面的牙形刺具有十分重要的意义。本文描述了采自新疆柯坪大湾沟和甘肃平凉官司庄剖面上奥陶统底界附近的牙形刺共13属18种。