Trilobite biofacies and sequence stratigraphy: an example from the Upper Ordovician of Oklahoma

There have been surprisingly few empirical investigations of the fundamental principle that the architecture of depositional sequences exerts considerable control on observed patterns of faunal distribution and replacement. In this paper, we examine trilobite associations in two sequences of the Upper Ordovician (Sandbian) Bromide Formation of southern Oklahoma. Cluster analysis and ordination of genus abundance data identified five lithofacies‐related biofacies that are also differentiated by diversity patterns. Biofacies of the transgressive system tract (TST) of successive sequences are more similar to each other than they are to biofacies in the highstand systems tract (HST) of the same sequence. This similarity likely records dominance of large, robust convex sclerites in taphonomically degraded samples from condensed, strongly winnowed grainstone and rudstone. Horizons with articulated exoskeletons of isoteline trilobites preserved by obrution deposits occur most commonly in the early HST and record behavioural aggregations. Grainstone and rudstone of the later HST are less winnowed than those of the TST and show less fragmentation and sorting of sclerites. These changes in taphonomic conditions preserve ecological patterns more clearly. In most biofacies, rarefied alpha diversity (samples) and gamma diversity (biofacies) of middle‐ and outer‐ramp HST deposits are greater than in the TSTs, and biofacies replace each other down ramp. Diversity patterns do not agree with model predictions and other data sets that indicate low beta and high alpha diversity in the TST, likely because of taphonomic degradation. Vertical replacement of biofacies is expressed by the appearance of peritidal facies in which trilobites are rare. Biofacies shifts also characterize sequence boundaries and are most profound in the inner‐ramp successions characterized by sharp facies offsets. Comparison with bathymetrically similar deposits in the Taconic foreland basin showed similar diversity trends along environmental gradients, with some differences in shallow‐water settings attributed to taphonomic differences.     

Trilobite faunal successions across the base of the Furongian Series in the Taebaek Group, Taebaeksan Basin, Korea

The base of the Furongian Series in the Sino-Korean Block has not been clearly defined due to the lack of the index taxon, Glyptagnostus reticulatus. The Sesong Formation of the Taebaek Group, Taebaeksan Basin, Korea, has been known to range from the Guzhangian Stage of the Cambrian Series 3 to the middle Furongian Series, hence embracing the base of the Furongian Series. Silicified polymerid trilobites were recovered from the middle part of the Sesong Formation. Described are a total of 18 polymerid species of 13 genera: Neodrepanura sp. 1, Teinistion sp. 1, Huzhuia sp. 1, Huzhuia sp. 2, Liostracina simesi, Liostracina sp. 1, Parachangshania monkei, Parachangshania rectangularis nov. sp., Placosema bigranulosum, Fenghuangella laevis nov. sp., Baikadamaspis jikdongensis nov. sp., Baikadamaspis sp. 1, Prochuangia mansuyi, Maladioides coreanicus, Alataspis sesongensis nov. gen., nov. sp., Chuangia sp. 1, and ceratopygids genus and species indeterminate 1 and 2. The stratigraphic occurrence of these trilobites provides a basis for recognition of five zones across the base of the Furongian Series (in ascending order): the Neodrepanura, Liostracina simesi, Fenghuangella laevis, Prochuangia mansuyi, and Chuangia zones. The Neodrepanura and Chuangia zones are provisionally adopted from the previous biostratigraphic scheme, while the three other ones are newly proposed. The recommended base of the Furongian Series in the Taebaek Group of Korea coincides with the base of the Fenghuangella laevis Zone, which appears to represent an episode of profound trilobite faunal turnover.     

The European Tertiary Neritiliidae (Mollusca, Gastropoda, Neritopsina): indicators of tropical submarine cave environments and freshwater faunas

The oldest freshwater neritiliid, Neritilia bisinuata , is described from the Middle Eocene of the Loire Basin. Another European species, N. neritinoides , ranging from the Lower Oligocene to Lower Miocene (Upper Burdigalian) is recognized; its habitat appears to have been freshwater, but very close to the sea. Two new marine neritiliid species from the Aquitaine Basin are described: Bourdieria favia sp. nov. from the Upper Oligocene and Pisulinella aucoini  sp. nov. from the Lower Miocene. A third undescribed species from the Lower Miocene is referred to the same family and related to Pisulinella . The Oligocene species has a strong spiral sculpture, a character completely absent in previously known neritiliid species. The genus Agapilia , founded on juvenile N. neritinoides and adult Vitta picta , appears to be a junior synonym of the genus Vitta. The associated occurrence of shells of the families Neritiliidae, Neritopsidae and Pickworthiidae (well-known inhabitants of Indo-West Pacific submarine caves) at Peyrère suggest the first occurrence of a characteristic assemblage of dark submarine caves during the Oligocene. Both factorial analysis and relative abundance show that at Peyrère these families are associated with other cryptic fossils (various gastropods, bivalves, Brachiopoda, corals, Annelida). However, there are indications of other submarine cave assemblages in various Cenozoic deposits from the Palaeocene to the middle Miocene.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 447–467.     

Quantitative palaeobiogeography: GIS, phylogenetic biogeographical analysis, and conservation insights

Aim  The utility of GIS-based and phylogenetic biogeographical analysis in palaeobiogeography is reviewed with reference to its ability to elucidate patterns of interest for modern conservation biology, specifically the long-term effects of invasive species.
Location  Emphasis is on biogeographical patterns in the Appalachian basin and mid-continent of North America during the Devonian. Global palaeobiogeographical patterns of the Cambrian are also considered.
Methods  Palaeobiogeographical patterns are assessed within a GIS framework, including both direct range reconstruction and niche modelling methods, and within phylogenetic biogeographical analysis. Biogeographical patterns are considered within multiple clades of fossil invertebrates, including trilobites, crustaceans, brachiopods, and bivalves.
Results  GIS-based analysis (including niche modelling methods) of Devonian invertebrates demonstrates a tightly correlated relationship between sea-level rises and range expansion, dispersal events, and species invasions. The predominance of range expansion and species invasions during the Late Devonian reduced opportunities for vicariant speciation during this interval. Comparison of phylogenetic biogeographical patterns between Cambrian and Devonian trilobites allows discernment of the relative roles of tectonics and eustacy in driving biogeographical patterns.
Main conclusions  GIS analysis and phylogenetic biogeography are powerful tools for analysing the coevolution of the Earth and its biota. Analyses can identify episodes of vicariance and geo-dispersal and produce testable hypotheses for further analysis within the fossil record.     

Facies architecture,depositional environments,and sequence stratigraphy of the Middle Cambrian Fasham and Deh-Sufiyan Formations in the central Alborz,Iran

Based on their lithologic characteristics and stratal geometries, the Middle Cambrian Fasham and Deh-Sufiyan Formations of the lower Mila Group in the Central Alborz, northern Iran, exhibit 39 lithofacies representing several supratidal to deep subtidal facies belts. The siliciclastic successions of the Fasham Formation are divided into two facies associations, suggesting deposition in a tide-dominated, open-mouthed estuarine setting. The mixed, predominantly carbonate successions of the Deh-Sufiyan Formation are grouped into ten facies associations. Four depositional zones are recognized on the Deh-Sufiyan ramp: basinal, outer ramp (deep subtidal associations), mid ramp (shallow subtidal to lower intertidal associations), and inner ramp (shoal and upper intertidal to supratidal associations). These facies associations are arranged in small-scale sedimentary cycles, i.e., peritidal, shallow subtidal, and deep subtidal cycles. These cycles reflect spatial differences in the reaction of the depositional system to small-scale relative sea-level changes. Small-scale cycles are stacked into medium-scale cycles that in turn are building blocks of large-scale cycles. Systematic changes in stacking pattern (cycle thickness, cycle type, and facies proportion) allow to reconstruct long-term changes in sea-level. Six large-scale cycles (S1–S6) have been identified and are interpreted as depositional sequences showing retrogradational (transgressive systems tract) and progradational (highstand systems tract) packages of facies associations. The six depositional sequences provide the basis for inter-regional sequence stratigraphic correlations and have been controlled by eustatic sea-level changes.     

Trilobite size-frequency distributions, recognition of instars, and phyletic size changes

Size-frequency analysis of over 5,000 Ordovician trilobites from the Teretiusculus Shales of the Builth inlier, central Wales, has revealed size distributions with counter intuitive shapes. Not only do most species show normal or slightly skewed distributions, despite the preponderance of moults, but there is no evidence of instar peaks. Such features can, however, be explained by reference to steady-state population structures of Recent marine arthropods, in which small individuals often form only a minor proportion of the post-larval population structure. Trilobite steady-state population structures would have differed in detail from species to species, but certain distribution shapes may have been characteristic of particular environments. These findings necessitate a reappraisal of previous work on trilobite size-frequency distributions, survivorship and recognition of instars. The Builth data also show the first clear evidence of phyletic size increase and parallel size changes in trilobites. ▭ Trilobites, size-frequency distributions, steady-state populations, instars, phyletic size changes.     

Carboniferous stratigraphy and depositional environments in the Ahnet Mouydir area (Algerian Sahara)

An up to 3,000-m-thick pile of Carboniferous rocks covers the northern fringe of the Precambrian Hoggar Massif (Touareg Shield) in southern Algeria, thus terminating the depositional history of the Palaeozoic in this sector of the North African Craton. The previous Devonian (Eifelian to Frasnian) palaeogeographic configuration of the area, characterized by ridges and shallow basins, is leveled by a widespread Famennian playa and lower Tournaisian delta sedimentation on a largely undifferentiated shelf. Tournaisian to Moscovian strata were deposited under open-marine, deltaic, shallow-subtidal, fluvial, and continental environments. The Carboniferous sequence can be subdivided into 12, largely interfingering, lithostratigraphic formations, which were dated by conodonts, ammonoids, foraminifers, and brachiopods, yielding a modified biostratigraphic framework of the area. The formations are stacked in four transgressive–regressive cycles, which include two major gaps, one during the middle Tournaisian, the other during the middle Visean to Serpukhovian. The oscillations of sea level can be traced into neighboring areas and seem coeval to early pulses of the Late Carboniferous/Early Permian glaciation of Gondwana. To a lesser degree they may reflect more local tectonic effects of the Variscan orogeny.     

Ordovician (Caradoc) Gastropoda of the Katkoyeh Formation, Kerman Province, Iran

Seven taxa of gastropods are described from the Ordovician (Caradoc) upper Katkoyeh Formation of the Kerman region, east-central Iran. Three are named species and four are under open nomenclature. The most abundant is the minute bellerophontiform Tritonophon peeli Horný, 1977, indicative of a shallow water plectonotid community. Shell material is not preserved in this species, but a small, simple bulbous protoconch is present. Two new species are Slehoferia pachyta and Nonorios kleistos, both with prominent thick-shelled conchs. Shell repair is documented in both of these forms, interpreted as a result of failed predation. Other micromorphic species occur in the samples, but only Tropidodiscus sp. and Nonorios? sp. are named. The Kerman assemblage is comparable directly with similar, closely coeval faunas in Bohemia, France, Portugal, Morocco, and Italy, corresponding with the Palaeozoic Mediterranean Province of northern peri-Gondwana.     

Trilobite body patterning and the evolution of arthropod tagmosis

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.     

Enters the shrew,some considerations on the Miocene palaeobiogeography of Iberian insectivores

The fossil record of the Iberian insectivores forms a subset of those found in central Europe. Comparison of the late early to early late Miocene record of the two areas shows that, particularly during the late Early Miocene, central European taxa have transient occurrences in Spain. Most taxa appear earlier and survive longer in central Europe. A notable exception is the gymnure Galerix, which extirpates earlier in central Europe, except for a transient occurrence in Germany just prior to its extinction. The main period of insectivore migrations is the late middle Miocene, although some of the taxa that enter remain restricted to the coastal areas. Overall, the pattern of distribution in time and space is best explained by the preference of insectivores for humid environments, as were found during the early Miocene and re-appeared at the end of the middle Miocene.     

The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks

The aim of this paper is to quantitatively investigate the spatial and temporal biogeographical relationships of the recovery of ammonoid faunas after the Permian-Triassic mass extinction using three complementary numerical approaches among which is a new, non-hierarchical clustering strategy. The faunal data set consists of a taxonomically homogenised compilation of the spatial and temporal occurrences of ammonoid genera within 20 Early Triassic Tethyan and Panthalassic sites ranging from 40°S to 70°N in palaeolatitudes. In addition to hierarchical cluster analysis (hCA) and nonmetric multidimensional scaling (NMDS), we introduce a third, new non-hierarchical clustering technique allowing the visualisation of a nonmetric interassemblages similarity structure as a connected network constructed without inferring additional internal nodes. The resulting network, which we call a “Bootstrapped Spanning Network” (BSN), allows the simultaneous identification of partially or totally nested as well as gradational linear or reticulated biogeographical structures.The identified interlocalities relationships indicate that the very beginning of the Early Triassic (Griesbachian) corresponds to a very simple biogeographical context, representing a time of great cosmopolitanism for ammonoids. This context shifts rapidly to a more complex configuration indicative of a more endemic and latitudinally-restricted distribution of the ammonoids during the middle and late Early Triassic (Smithian and Spathian). From an evolutionary dynamic point of view, our results illustrate a very rapid (less than ca. 1.4 myr) Early Triassic recovery of the ammonoid faunas, in contrast to many other marine organisms. This recovery is linked with a marked increase in the overall biogeographical heterogeneity, and parallels the formation of a latitudinal gradient of taxonomic richness, which may be essentially controlled by the progressive intensification of the gradient of sea surface temperature. From a methodological point of view, we show that a BSN is a simple, intuitively legible picture of the nested as well as gradational taxonomic similarity relationships, hence providing a good synthesis (and additional insights) between hierarchical clustering and ordination in reduced space results.     

Biostratigraphy and palaeobiogeography of the Cambrian genusHipponicharion (Ostracoda)

The bradoriid genusHipponicharion Matthew, 1886 is one of the oldest known Cambrian ostracods. Until now, five species have been documented and another three taxa have been tentatively assigned to one or another species.Hipponicharion has been recorded from Poland, Germany, Morocco as well as from Canada and Great Britain.Hipponicharion hispanicum n. sp. represents the first Cambrian ostracod documented from Spain.Hipponicharion taidaltensis n. sp., formerlyH. aff.geyeri Hinz-Schallreuter, 1993 andH. elickii n. sp., formerlyH. cf.eos sensuElicki, 1994 have also been restudied in this paper.Hipponicharion seems to be restricted to the Acadobaltic Province sensuSdzuy (1972). The systematic relationships of the Bradoriida, which is controverse, is briefly outlined, emphasizing close affinities to the Ostracoda.      

Genus age,provincial area and the taxonomic structure of marine faunas

Species are unevenly distributed among genera within clades and regions, with most genera species-poor and few species-rich. At regional scales, this structure to taxonomic diversity is generated via speciation, extinction and geographical range dynamics. Here, we use a global database of extant marine bivalves to characterize the taxonomic structure of climate zones and provinces. Our analyses reveal a general, Zipf–Mandelbrot form to the distribution of species among genera, with faunas from similar climate zones exhibiting similar taxonomic structure. Provinces that contain older taxa and/or encompass larger areas are expected to be more species-rich. Although both median genus age and provincial area correlate with measures of taxonomic structure, these relationships are interdependent, nonlinear and driven primarily by contrasts between tropical and extra-tropical faunas. Provincial area and taxonomic structure are largely decoupled within climate zones. Counter to the expectation that genus age and species richness should positively covary, diverse and highly structured provincial faunas are dominated by young genera. The marked differences between tropical and temperate faunas suggest strong spatial variation in evolutionary rates and invasion frequencies. Such variation contradicts biogeographic models that scale taxonomic diversity to geographical area.     

Araucariaceae (Pinopsida): Aspects in palaeobiogeography and palaeobiodiversity in the Mesozoic

The paper examines recent information on the history of the ‘southern hemisphere’ conifer family Araucariaceae and gives supplementary notes to previously published monographs. Important data from the Mesozoic fossil record are presented and summarized to describe the ancient diversity and distribution of Araucariaceae. Information on the origin of the family and oldest fossil records of the present-day genera are assembled. Existing models on phylogenetic relationships within the family inferred from molecular data are compared with relationships between the genera determined from morphological data. Reasons for disappearance of representatives of this conifer family in the latest Cretaceous in North America and Europe are briefly discussed.     

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.     

Systematics,palaeoecology and palaeobiogeography of the Neogene fossil sharks from the Azores (Northeast Atlantic)

The Azores Archipelago is a group of isolated islands located in the North Atlantic Ocean. One of these oceanic islands – Santa Maria – exhibits marine fossiliferous sediments of late Miocene/early Pliocene and also of Pleistocene age. Recent research provided new selachian fossil material, with three new records (Carcharias acutissima, Megaselachus megalodon, and Carcharhinus cf. leucas) increasing the number of fossil sharks reported from the Azores (Santa Maria Island) to seven species (Notorynchus primigenius, C. acutissima, Cosmopolitodus hastalis, Paratodus benedenii, Isurus oxyrinchus, M. megalodon, and C. cf. leucas). So far, no teeth of batoids or small sharks have been found despite the screen-washing of several sediment samples from Santa Maria. The Azorean Mio-Pliocene selachian fauna clearly differs from those described from sediments deposited on continental shelves, in which batoids and small benthic sharks (e.g., scyliorhinids) are usually well represented. During the late Miocene/early Pliocene, subtropical to warm-temperate seas were prevalent in the area of the Azores, as deduced from palaeontological, geological and isotopic data, all indicating a warmer climate than in the present.     

Continental gateways and the dynamics of mammalian faunas

Continental gateways occur where mountainous topography interacts with changing climate and sea level to open or close dispersal corridors. The interaction of permeable or impermeable montane barriers with changing or stable climate yields four biogeographic states, each associated with changes in diversification rates and ecological structure of faunas. For example, permeable montane barriers and climatic stability result in low rates of immigration and extinction, elevated endemic speciation, and stable ecological structure. Three examples from the mammalian fossil record test these scenarios. (1) In Miocene faunas of Pakistan, immigration rates peaked and faunal proportions changed during an interval of cooling and open corridors. (2) In Miocene faunas of Spain, elevated extinction and origination rates and changing trophic structure occurred during regional aridification with open corridors. (3) In Quaternary faunas of South Africa, ungulates experienced range reductions and elevated extinction during the transition from glacial to interglacial climates as corridors closed.