The Origin and Initial Rise of Pelagic Cephalopods in the Ordovician

Background

During the Ordovician the global diversity increased dramatically at family, genus and species levels. Partially the diversification is explained by an increased nutrient, and phytoplankton availability in the open water. Cephalopods are among the top predators of todays open oceans. Their Ordovician occurrences, diversity evolution and abundance pattern potentially provides information on the evolution of the pelagic food chain.

Methodology/Principal Findings

We reconstructed the cephalopod departure from originally exclusively neritic habitats into the pelagic zone by the compilation of occurrence data in offshore paleoenvironments from the Paleobiology Database, and from own data, by evidence of the functional morphology, and the taphonomy of selected cephalopod faunas. The occurrence data show, that cephalopod associations in offshore depositional settings and black shales are characterized by a specific composition, often dominated by orthocerids and lituitids. The siphuncle and conch form of these cephalopods indicate a dominant lifestyle as pelagic, vertical migrants. The frequency distribution of conch sizes and the pattern of epibionts indicate an autochthonous origin of the majority of orthocerid and lituitid shells. The consistent concentration of these cephalopods in deep subtidal sediments, starting from the middle Tremadocian indicates the occupation of the pelagic zone early in the Early Ordovician and a subsequent diversification which peaked during the Darriwilian.

Conclusions/Significance

The exploitation of the pelagic realm started synchronously in several independent invertebrate clades during the latest Cambrian to Middle Ordovician. The initial rise and diversification of pelagic cephalopods during the Early and Middle Ordovician indicates the establishment of a pelagic food chain sustainable enough for the development of a diverse fauna of large predators. The earliest pelagic cephalopods were slowly swimming vertical migrants. The appearance and early diversification of pelagic cephalopods is interpreted as a consequence of the increased food availability in the open water since the latest Cambrian.     

The Bohemo‐Iberian regional chronostratigraphical scale for the Ordovician System and palaeontological correlations within South Gondwana

The Bohemo‐Iberian regional scale for South Gondwana, involving the ‘Mediterranean Province’, comprises five regional stages (Arenigian, Oretanian, Dobrotivian, Berounian and Kralodvorian) plus the global Tremadocian and Hirnantian. The predominance of shallow‐water taxa in those high‐latitude faunas imposes serious difficulties for correlating the regional succession with the formal global chronostratigraphy because of the almost total absence of the key graptolites and conodonts defining the base of the standard series, stages and stage slices. Instead, the abundant benthic faunas (trilobites, brachiopods, molluscs, echinoderms) of South Gondwanan origin largely dominated in the area from the middle Darriwilian to the late Katian. The poleward faunal migration of originally Avalonian, Baltic, Laurentian and even Asiatic taxa during the Boda Event of global warming progressively ends with the endemicity in the region, where the ensuing benthic assemblages were severely affected by the Hirnantian glaciation. The regional scale significantly improves the precision of correlations between Ordovician strata from SW and central Europe, North Africa and a large part of the Middle East. An updated record of palaeontological data from areas where Mediterranean faunas remain practically unknown, or are still poorly investigated, is also included. Palaeobiogeographical relationships based on the distribution of faunas across South Gondwana are suggested as an improvement for positioning many territories in modern palaeogeographical reconstructions and offer a constructive approach to problems related to the pre‐Variscan and pre‐Alpine orogenic puzzles.     

BRYOZOAN MUD-MOUNDS FROM THE UPPER ORDOVICIAN JIFARAH (DJEFFARA) FORMATION OF TRIPOLITANIA, NORTH-WEST LIBYA

Abstract:  A bryozoan fauna from carbonate mud-mounds is described from subsurface well cores from the Upper Ordovician (Lower Ashgill) Jifarah (Djeffara) Formation of Tripolitania, north-west Libya. Among a diverse assemblage dominated by trepostomes, nine species of bryozoans are identified, including Jifarahpora libyensis gen. et sp. nov. Delicate and robust branching, encrusting and nodular bryozoan growth forms are all common. The bryozoan-rich limestones are mostly mudstones and wackestones, with bafflestone and floatstone textures, but the mounds apparently lack organic framework and microbial fabrics. Regional geophysical data indicate rapid thickness changes between wells, where mound complexes locally up to 100 m thick had limited topographic relief over the surrounding sea floor. The mounds formed in a high-latitude, cool-water carbonate belt that extended widely across the northern margin of Gondwana. Quaternary analogues from the Great Australian Bight suggest that these early Ashgill mounds may have developed in slope environments during an episode of glacial lowstand that preceded the late Ashgill, Hirnantian glacial event.     

Septal fracture in Nautilus: implications for cephalopod paleobathymetry

Analysis of septal geometry, as embodied in the septal strength index model developed by Westermann (1973, Lethaia 6). , has become a prime avenue for estimating living depths of fossil cephalopods. We have examined the fracturing of Nautilus septa, and its bearing on strength index, by inducing septal rupture under the action of hydrostatic pressure. We found that: (1) septa which actually fail under pressure are not generally weakest as defined by their strength indices; (2) septal strength as defined by the strength index is not correlated with rupture pressure; and (3) most instances of septal failure originate in septal sutures, not in the septa. These results indicate that: (1) septal strength index does not yield wholly reliable strength or depth estimates; and (2) the shortcomings of the strength index model stem from its inability to account for complexities of mechanical failure in morphologically complex cephalopod shells.     

Pelagic cephalopods of the central Mediterranean Sea determined by the analysis of the stomach content of large fish predators

The pelagic cephalopod fauna of the central Mediterranean Sea was investigated through stomach content analyses of large fish predators. A total of 124 Xiphias gladius, 22 Thunnus thynnus, 100 Thunnus alalunga, and 25 Tetrapturus belone were analyzed. Overall, 3,096 cephalopods belonging to 23 species and 16 families were identified. The cephalopod fauna in the study area is dominated by Sepiolidae, Ommastrephidae, and Onychoteuthidae. The sepiolid Heteroteuthis dispar was the most abundant species (n?=?1,402) while the ommastrephid Todarodes sagittatus showed the highest biomass. They can be considered key-species in the pelagic food web of the study area. The neutrally buoyant Histioteuthis bonnellii, H. reversa, and Chiroteuthis veranyi seem to characterize the deeper water layers. Given the difficulty in sampling pelagic cephalopods, the presence of cephalopod beaks in the stomach of predators represents a fundamental tool to assess the biodiversity and the ecological importance of these taxa in the marine ecosystem.     

Late Ordovician brachiopods from eastern North Greenland: equatorial offshore migration of the Red River fauna

Abstract: Late Ordovician rhynchonelliformean brachiopods, typical of the North American Red River fauna, are found sporadically in the Børglum River Formation of the Centrum Sø area, Kronprins Christian Land, eastern North Greenland. The geographical distribution of this characteristic brachiopod fauna is thus extended to the easternmost extremity of the Laurentian craton. The assemblage compares specifically with the Hiscobeccus brachiopod fauna, based on key taxa such as notably Hiscobeccus gigas (Wang, 1949), and indicates a late Katian age for this part of the succession. For the first time, this typically inland, shallow‐water fauna is found associated with genera like Bimuria, suggesting a transitional marginal facies with outer shelf benthos. The current study describes a Hiscobeccus fauna that lived at the seaward edge of its preferred habitat. Furthermore, an unpublished Hiscobeccus fauna, from the Børglum River Formation of Peary Land, central North Greenland, as well as several occurrences from the Kap Jackson and Cape Calhoun formations in various parts of Washington Land, western North Greenland, are described here. These show a distinct shift from older strata containing H. capax (Conrad, 1842) to younger strata exclusively yielding specimens of H. gigas. As H. gigas occurs in the upper part of the Cape Calhoun Formation in Washington Land, it indicates that the upper boundary of the Cape Calhoun Formation is considerably younger than previous estimates, reaching into the uppermost Katian (middle Cautleyan–Rawtheyan). The Cape Calhoun Formation correlates with the upper member of the Børglum River Formation and further demonstrates that the Hiscobeccus fauna was widespread in Laurentian marginal settings of North Greenland. Even though the Hiscobeccus fauna was pan‐continental during the late Katian (Richmondian), it possesses a strong provincial signal during the later Ordovician. The new occurrences indicate that this fauna extended to the north‐eastern margin of the Laurentian Craton. It lived in close association with cosmopolitan faunal elements that may have been the earliest sign of the succeeding invasion of migrants from Baltica that arrived later during the Hirnantian. The offshore migration of this atypical Hiscobeccus fauna likely demonstrates the path of warm‐water currents as the Centrum Sø locality was located at the equator during the Late Ordovician.     

Middle Ordovician cephalopod biofacies and palaeoenvironments of Baltoscandia

During the Middle Ordovician cephalopods became an important part of the macrofauna of the Baltoscandian carbonate platform. The earliest cephalopod abundance peak was reached during the early Darriwilian, within the Kunda Stage Yangtzeplacognathus crassus and Lenodus pseudoplanus conodont zones. In sediments of this time interval large orthoconic cephalopods often occur in masses with more than one specimen per square‐meter on bedding surfaces. The assemblages are characterized by the strong dominance of often large endocerids. In proximal depositional settings coiled tarphycerids and other cephalopod groups are an important additional component. In the most distal settings orthocerids are the most important secondary component. Correspondence Analysis of assemblages throughout Baltoscandia revealed three distinct biofacies, which here are termed Orthocerid, Proterovaginoceras and Anthoceras Biofacies, respectively. The biofacies reflect differences in depth and proximity to the shoreline and are consistent with the Baltoscandian Confacies Belts. Spatial changes in absolute abundance and taxonomic composition indicate increased original cephalopod population densities and habitat expansion within the Y. crassus and L. pseudoplanus conodont zones. A nearly coeval abundance peak in a similar facies in South China indicates supraregional causes of the mass occurrence, probably reflecting a globally increased nutrient availability in the water column during the Darriwilian.     

Pelagic cephalopods from eastern Australia: species composition, horizontal and vertical distribution determined from the diets of pelagic fishes

The distribution and relative biomass of cephalopods from pelagic waters off eastern Australia was examined between 1997 and 2004 from stomach contents of swordfish, yellowfin tuna and dolphinfish taken in the domestic longline fishery. A total of 38 taxa from 19 families were identified. Comparison of the species composition of the three predators indicated pronounced differences in cephalopod species composition. In swordfish, species of the family Ommastrephidae, particularly Ommastrephes bartramii (Lesueur 1821) and Nototodarus gouldi (McCoy 1888) dominated, whereas a more diverse mix of species was identified from yellowfin-sampled cephalopods. Todaropsis eblanae (Ball 1841) was the main cephalopod sampled from the surface-dwelling dolphinfish. For swordfish-sampled cephalopods, significant relationships were found between biomass and season, fluorescence and year. In yellowfin tuna, cephalopod biomass was significantly correlated with season, area and sea surface temperature. Significant factors differed between predator-sampler, possibly reflecting the limits of the predator, but could also give insights into individual cephalopod species distributions. However, the increase in cephalopod biomass over summer in both swordfish and yellowfin tuna suggested cephalopod biomass was higher over summer in the region.     

The La Voulte-sur-Rhône Lagerstätte (Middle Jurassic,France)

The La Voulte Lagerstätte (Jurassic, France) is characterized by a diverse and exceptionally well-preserved fauna. It was located along the western Tethyan margin where the submarine relief was steep. The sedimentation was dominated by marls suggesting a low energy and deep-water depositional environment. The La Voulte biota is remarkable for its biodiversity with about 60 different species. The major components are the arthropods (50% of the species), the cephalopods (10%) and the echinoderms (10%). Among the arthropods, the composition and some anatomical features suggest a relatively deep-water environment characterized by dim-light conditions. The cephalopods and other organisms (pycnogonids, asterids), have extant analogues that all live in deep-water niches always exceeding 200 m. The fauna as a whole bears the characteristic features of the present-day bathyal fauna. The La Voulte Lagerstätte may be one of the rare Jurassic Lagerstätten, if not the unique, to have fossilized a deep marine fauna.     

Renal organs of cephalopods: a habitat for dicyemids and chromidinids

The renal organs of 32 species of cephalopods (renal appendage of all cephalopods, and renal and pancreatic appendages in decapods) were examined for parasite fauna and for histological comparison. Two phylogenetically distant organisms, dicyemid mesozoans and chromidinid ciliates, were found in 20 cephalopod species. Most benthic cephalopods (octopus and cuttlefish) were infected with dicyemids. Two pelagic cephalopod species, Sepioteuthis lessoniana and Todarodes pacificus, also harbored dicyemids. Chromidinid ciliates were found only in decapods (squid and cuttlefish). One dicyemid species was found in branchial heart appendages of Rossia pacifica. Dicyemids and chromidinids occasionally occurred simultaneously in Euprymna morsei, Sepia kobiensis, S. peterseni, and T. pacificus. The small-sized cephalopod species, Idiosepius paradoxus and Octopus parvus, harbored no parasites. Comparative histology revealed that the external surface of renal organs varies morphologically in various cephalopod species. The small-sized cephalopod species have a simple external surface. In contrast, the medium- to large-sized cephalopod species have a complex external surface. In the medium- to large-sized cephalopod species, their juveniles have a simple external surface of the renal organs. The external surface subsequently becomes complicated as they grow. Dicyemids and chromidinids attach their heads to epithelia or insert their heads into folds of renal appendages, pancreatic appendages, and branchial heart appendages. The rugged and convoluted external surface provides a foothold for dicyemids and chromidinids with a conical head. They apparently do not harm these tissues of their host cephalopods.     

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.     

A cephalopod fauna from the middle Ordovician of Saudi Arabia

A small collection of cephalopods from the middle Ordovician of the Tabuk region of Saudi Arabia consists entirely of orthoconic longicones tentatively referred to the actinocerid genus Mesaktoceras? and pseudorthocerid Sactoceras?. Both taxa have Australasian affinities. Circumpolar Gondwana cephalopod faunas in the mid Ordovician were of low diversity and dominated by orthocerids, whilst actinocerids were largely restricted to low palaeolatitudes. The presence of Mesaktoceras? and Sactoceras? in a benthonic fauna belonging to the Neseuretus community is regarded as anomalous. Comparison is made with the occurrence of Trocholites in middle Ordovician Iberian successions. It has been suggested that Trocholites may have arrived in the Iberian area from Baltica through the transfer of bodies of warm water in storm masses. Mesaktoceras? and Sactoceras? may have arrived (albeit temporarily) in Saudi Arabia from Australasia through a similar process.     

Ecological biogeography of cephalopod molluscs in the Atlantic Ocean: historical and contemporary causes of coastal diversity patterns

Aim One of the most recognized ecological paradigms on earth is the increase in species richness from the poles towards the equator. Here we undertake a comprehensive survey of the latitudinal gradients of species richness (LGSR) of coastal cephalopod fauna in the western (WA) and eastern margins (EA) of the Atlantic Ocean, and test climate and non‐climate theories to explain the variation in diversity. Location The coastal Atlantic Ocean. Methods The diversity and geographical ranges of coastal cephalopods were investigated by means of an exhaustive survey of the primary literature, reports and on‐line data bases. In order to test the productivity, ambient energy and area hypotheses, we investigated the relationship between diversity and net primary production (NPP), sea surface temperature (SST; measure of solar energy input) and continental shelf area, respectively. Results LGSR of cephalopod molluscs are present at both Atlantic coasts, but are quite distinct from each other. Historical processes (rise of the Central American Isthmus, formation of ‘Mare Lago’ and glaciations) explained much of the shape and the zenith of LGSR. Contemporary climate and non‐climate variables also each explained over 83% and 50% of the richness variation in WA and EA, respectively, and the best fitted models accounted for > 92% of the variance. By combining latitude with depth a strong Rapoport effect was observed in WA but not in EA. Main conclusions Besides the evolutionary history, we demonstrate that the contemporary environmental gradients (SST and NPP), shelf area and extent of coral habitat can predict many of the diversity patterns. The longitudinal difference in Rapoport's bathymetric rule is attributed to western fauna specialization to shallow coral reef habitats and greater ecological tolerance of eastern fauna to upwelling ecosystem dynamics. A combined approach of historical biogeography and species–area–energy theories was essential to fully understand broad‐scale variation in cephalopod biodiversity.     

Approaches to resolving cephalopod movement and migration patterns

Cephalopod movement occurs during all phases of the life history, with the abundance and location of cephalopod populations strongly influenced by the prevalence and scale of their movements. Environmental parameters, such as sea temperature and oceanographic processes, have a large influence on movement at the various life cycle stages, particularly those of oceanic squid. Tag recapture studies are the most common way of directly examining cephalopod movement, particularly in species which are heavily fished. Electronic tags, however, are being more commonly used to track cephalopods, providing detailed small- and large-scale movement information. Chemical tagging of paralarvae through maternal transfer may prove to be a viable technique for tracking this little understood cephalopod life stage, as large numbers of individuals could be tagged at once. Numerous indirect methods can also be used to examine cephalopod movement, such as chemical analyses of the elemental and/or isotopic signatures of cephalopod hard parts, with growing interest in utilising these techniques for elucidating migration pathways, as is commonly done for fish. Geographic differences in parasite fauna have also been used to indirectly provide movement information, however, explicit movement studies require detailed information on parasite-host specificity and parasite geographic distribution, which is yet to be determined for cephalopods. Molecular genetics offers a powerful approach to estimating realised effective migration rates among populations, and continuing developments in markers and analytical techniques hold the promise of more detailed identification of migrants. To date genetic studies indicate that migration in squids is extensive but can be blocked by major oceanographic features, and in cuttlefish and octopus migration is more locally restricted than predictions from life history parameters would suggest. Satellite data showing the location of fishing lights have been increasingly used to examine the movement of squid fishing vessels, as a proxy for monitoring the movement of the squid populations themselves, allowing for the remote monitoring of oceanic species.     

The bearing of the new Late Cambrian monoplacophoran genus Knightoconus upon the origin of the Cephalopoda

Yochelson, E. L., Flower, R. H. & Webers, G. F.: The hearing of the new Late Cambrian monoplacophoran genus Knightoconus upon the origin of the Cephalopoda.
Knightoconus , a new genus of the Hypseloconidae (Mollusca: Monoplacophora) from rocks of early Franconian age in Antarctica, is multiseptate. The multiple septa are a criticàl feature to be expected in a form ancestral to cephalopods. Fossil cephalopods, however, invariably have a siphuncle as well as septa; some gastropods, some hyolithids, and some monoplacophorans also have septa but lack a siphuncle. Therefore, only the siphuncle can be considered a unique and particularly significant feature of the cephalopod shell. Hypothetical reconstructions of molluscan anatomy support the notion that cephalopods may have been derived directly from a hypseloconid having a high, slightly curved, multiseptate, bilaterally symmetrical shell, by the subsequent development of a siphuncle.     

Oldest known Dicoelosia and Epitomyonia,deep water brachiopods from the Beiguoshan Formation (Middle Katian,Upper Ordovician), Shaanxi,north China

Abstract: A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender‐lobed form with a concavoconvex profile to the strongly equibiconvex, fat‐lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

The relationships between Late Ordovician sea-level changes and faunal turnover in western Baltica: Geochemical evidence of oxic and dysoxic bottom-water conditions

The late Sandbian to the early Katian (Late Ordovician) in southeastern Norway is dominated by marine mudstones that contain an exceptional spectrum of macrofauna and trace element geochemistry, recording both abrupt and gradual faunal changes during major sea-level and environmental shifts. This study investigates the relationships between sea-level changes and their influence on the source of clastic material, oxygen levels in the bottom water and faunal changes, with special emphasis on the brachiopod fauna. Trace element ratios indicate that nearly stable upper dysoxic bottom-water conditions prevailed in the northwestern part of the Baltoscandian Sea. The exception is during the major shallowing in the earliest Katian, when there was an abrupt shift to oxic conditions as the sea bottom came within a normal storm wave base. Nonetheless, the pre-shallowing epibenthic fauna (though not the shelly endobenthic) in this area was rich and diverse and two major immigration phases of new brachiopod taxa are seen well before the shallowing. This indicates that the immigration of new taxa was not the result of an increase in oxygen content in the bottom water. More brachiopod genera stayed during the abrupt shallowing and increased oxic level than did during the following gradual return to deeper, dysoxic environments. The major brachiopod immigration phases took place markedly earlier in this northwestern part of the Baltoscandian Sea than in the central part (Eastern Baltic) and possibly also the comparable faunal turnover in Laurentia. The following disappearance of taxa during an early Katian transgressive event coincided with the faunal turnover in the shallow-water environments of the East Baltic.The depositional history was consistent within the investigated mudstone-dominated offshore facies of the Oslo Region, which are comparable to the Central Baltoscandian Confacies Belt of Baltoscandia. The composition of the siliciclastic material which was deposited in the basin was nearly constant through the sequence, though the land area expanded to include ophiolites, expressed by Cr enrichment, coinciding with the major sea-level drop. The Cr enrichment indicates that, by that time at least, the ophiolite complexes north of the Oslo Region became subaerially exposed. This enrichment thus forms a potential geochemical marker horizon representing the basal Katian Stage.