Population dynamics of galeate acritarchs at the Cambrian–Ordovician transition in the Algerian Sahara

Galeate acritarchs are a major component of Late Cambrian and Early Ordovician palynological assemblages. The populations of galeate acritarchs from the Cambrian–Ordovician transition section in borehole Nl-2 of the Algerian Sahara show a gradual increase in process length and in complexity of the process structures through the succession. While the number of striate elements at the process base, the number of distal ramifications, and the number of membranes between the processes increases progressively, the central body diameter shows only minor variation within the succession. The galeate acritarchs are here interpreted as probably being resting cysts of a microorganism similar to modern dinoflagellates. Published laboratory culture experiments on living dinoflagellates document a considerable morphological variation with respect to the process morphology of the resting cysts that can be produced by a single biologically defined species. Based on these experiments, it is possible to interpret the morphological changes in the galeate acritarchs as being possibly related to changing ecological conditions. By analogy to the cyst distribution of modern dinoflagellates, changing salinity may have played an important role. Depending on environmental parameters, and perhaps on the maturity attained before rupture of the outer membrane during cyst formation, the galeate acritarchs may show a wide variety of process morphologies that have to date been used to describe four genera (morphogenera) and 84 species (morphospecies).     

A nearshore–offshore trend in acritarch distribution from the Early–Middle Ordovician of the Yangtze Platform, South China

The stratigraphical interval of the late Early Ordovician Didymograptus deflexus and the early Middle Ordovician Azygograptus suecicus graptolite Biozones was investigated from seven sections from the upper Yangtze Platform, southern China. These are located on different parts of the platform, between the nearshore environments of the Kunming area, Yunnan Province, and the offshore carbonate shelf of the Yichang area, Hubei Province. The assemblages recovered from the different parts of the platform vary both in terms of diversity and composition. The nearshore environments show low diversity assemblages with about 10 acritarch species, whereas the offshore shelf environments reflect higher diversities with about 40 species. The composition of the assemblages also changes from simple morphologies (micrhystrids, leiosphaerids, fusiform acritarchs) in nearshore environments to specimens with longer and more complexly branched processes on the shelf. The polygonomorph acritarchs are common over all the upper Yantze Platform, while the acanthomorph genera Baltisphaeridium and Peteinosphaeridium are the most abundant taxa on the offshore carbonate shelf area. Phylogenetic or sea-level changes are probably not responsible for the compositional and diversity changes that occurred during the investigated interval. This study confirms previous interpretations that poorly diversified Palaeozoic acritarch assemblages occur in neritic environments and more complex, highly diversified assemblages are found on the shelf.     

Activity patterns in the Sahara Desert: an interpretation based on cross-sectional geometric properties

The Garamantian civilization flourished in modern Fezzan, Libya, between 900 BC and 500 AD, during which the aridification of the Sahara was well established. Study of the archaeological remains suggests a population successful at coping with a harsh environment of high and fluctuating temperatures and reduced water and food resources. This study explores the activity patterns of the Garamantes by means of cross-sectional geometric properties. Long bone diaphyseal shape and rigidity are compared between the Garamantes and populations from Egypt and Sudan, namely from the sites of Kerma, el-Badari, and Jebel Moya, to determine whether the Garamantian daily activities were more strenuous than those of other North African populations. Moreover, sexual dimorphism and bilateral asymmetry are assessed at an intra- and inter-population level. The inter-population comparisons showed the Garamantes not to be more robust than the comparative populations, suggesting that the daily Garamantian activities necessary for survival in the Sahara Desert did not generally impose greater loads than those of other North African populations. Sexual dimorphism and bilateral asymmetry in almost all geometric properties of the long limbs were comparatively low among the Garamantes. Only the lower limbs were significantly stronger among males than females, possibly due to higher levels of mobility associated with herding. The lack of systematic bilateral asymmetry in cross-sectional geometric properties may relate to the involvement of the population in bilaterally intensive activities or the lack of regular repetition of unilateral activities.     

Trace fossils on a Late Ordovician glacially striated pavement in Algeria

An exceptional exposure of Late Ordovician glaciogenic sediments crops out in Dider, SE Algeria, within the Tassili N'Ajjer region. The sediments consist of sandstones and diamictites sandwiched between a Mid Ordovician fluvial and tidal sandstone (In Tahouite Formation) below and Early Silurian shale (Oued Imirhou Formation) above. Stratigraphic discontinuities within the Late Ordovician glaciogene succession include palaeovalley incisions and glacially striated pavements. Striation and fluting of a soft-bedded sediment beneath an ancient ice sheet is supported by abundant dewatering structures and soft-sediment gouges interpreted to have been produced by the action of stone ploughing. In Dider, two types of previously undescribed circular structures sit in negative relief on this glacial pavement, namely 1) paired thumb-shaped impressions 2 cm in diameter and 3 mm in depth, and 2) a 5 cm wide impression with 3 segmented nested cycles. A framboid or aggregate origin may be appropriate for the smaller of the features but the larger impression is interpreted as biogenic: internal complexity is characteristic, discounting concretion moulds and water escape structures. A biogenic origin as a coelenterate resting trace is proposed with speculation on conditions of exceptional preservation in an ancient periglacial environment.     

Cryptospore Assemblages from the Ordovician/silurian Boundary in the Puna Region, North-west Argentina

Palynomorph assemblages have been recovered from deposits believed to straddle the Ordovician/Silurian boundary, from the upper member of the Salar del Rincón Formation, in the Puna region of north-west Argentina. The palynomorph assemblages are dominated by terrestrial cryptospores, but also contain marine elements (acritarchs, prasinophycean algae and chitinozoans). The cryptospore assemblages are similar in composition to those described from coeval deposits worldwide, suggesting that the producers were cosmopolitan and tolerated a wide range of climatic conditions. They are correlated with the Imperfectotriletes spp. Interval Biozone (sub-biozone α) of the Imperfectotriletes spp.- Laevolancis divellomedia Assemblage Biozone, indicating a Hirnantian (latest Ordovician) age. Acritarchs include late Ordovician species such as Eupoikilosusa striata and Villosacapsula cf. setosapellicula , that coexist with the Llandovery species Dactylofusa estillis. Based on lithological and palynological evidence, an early Llandovery, or a late Hirnantian (post-glacial) age is proposed for the basal part of the upper member of the Salar del Rincón Formation.     

Palaeoenvironmental significance of cool‐water microbialites in the Darriwilian (Middle Ordovician) of Sweden

Well‐developed oncoids and centimetre‐sized stromatolites are reported for the first time from the Darriwilian (Middle Ordovician) cool‐water ‘orthoceratite limestone’ at Kinnekulle, Västergötland, Sweden. The characteristics and stratigraphical distribution of these microbialites show an apparent relationship to fluctuations in relative sea level. The most abundant and well‐developed oncoids occur in stratigraphical intervals that are characterized by notable sea‐level lowstands. Stromatolites, which share many compositional characteristics with the oncoids, are essentially confined to a single bed associated with an especially prominent lowstand. Stromatolite‐like lamination also occurs in the uppermost part of the studied succession, but this feature may be of abiogenic origin. The microbialites appear to be originally calcareous, but synsedimentary iron‐ and/or phosphate‐enriched laminae are conspicuous, and secondary substitution by coarse calcite and barite is common. Iron staining is most prominent in poorly preserved specimens. Diagenesis has occluded the identity of the producers of these microbialites, but characteristics of associated endolithic borings suggest that they were formed in photic waters. The laminated fabrics of the documented microbialites record a depositional environment sensitive to high‐frequency environmental change. Most significantly, the microbialites have provided important information about the depositional environment of their enigmatic host limestone, and the collective observations challenge the notion that the studied strata were deposited in a deep shelf to basinal environment – rather, it appears that they are to a large extent, shallow‐water deposits, formed in waters only a few tens of metres deep.     

The presence–absence matrix reloaded: the use and interpretation of range–diversity plots

Aim A great deal of information on distribution and diversity can be extracted from presence–absence matrices (PAMs), the basic analytical tool of many biogeographic studies. This paper presents numerical procedures that allow the analysis of such information by taking advantage of mathematical relationships within PAMs. In particular, we show how range–diversity (RD) plots summarize much of the information contained in the matrices by the simultaneous depiction of data on distribution and diversity. Innovation We use matrix algebra to extract and process data from PAMs. Information on the distribution of species and on species richness of sites is computed using the traditional R (by rows) and Q (by columns) procedures, as well as the new Rq (by rows, considering the structure of columns) and Qr (by columns, considering the structure by rows) methods. Matrix notation is particularly suitable for summarizing complex calculations using PAMs, and the associated algebra allows the implementation of efficient computational programs. We show how information on distribution and species richness can be depicted simultaneously in RD plots, allowing a direct examination of the relationship between those two aspects of diversity. We explore the properties of RD plots with a simple example, and use null models to show that while parameters of central tendency are not affected by randomization, the dispersion of points in RD plots does change, showing the significance of patterns of co‐occurrence of species and of similarity among sites. Main conclusion Species richness and range size are both valid measures of diversity that can be analysed simultaneously with RD plots. A full analysis of a system requires measures of central tendency and dispersion for both distribution and species richness.     

The earliest myodocopes: ostracodes from the late Ordovician Soom Shale Lagerstätte of South Africa

The late Ordovician Soom Shale Lagerstätte of South Africa has yielded Myodoprimigenia fistuca n. gen. and n. sp., the earliest and only known Ordovician occurrence of myodocopes, one of the major groups of ostracodes. M. fistuca is a likely sister group of the Upper Silurian 'cypridinid' myodocopes and allied forms. It had a thin, lightly mineralized and flexible shell with microstructures resulting from in vivo calcification processes. It probably fed on cephalopod carrion, thus extending evidence for a carnivorous scavenging lifestyle in ostracodes back by 200  Ma. The species was probably nektobenthic and thus consistent with the notion that the origin of the late Silurian pelagic myodocopes - and therefore of pelagic ostracodes - is to be charted in a benthic to pelagic ecological shift in the group.     

Evenness and diversity in Upper Cambrian – Lower Ordovician trilobite communities from the Central Andean Basin (Cordillera Oriental,Argentina)

Community evenness has recently received much attention, either because it is related to ecosystem functioning or because it may affect estimation of diversity. Temporal and environmental trends in diversity and evenness of trilobite communities during the Late Cambrian – Early Ordovician of the Cordillera Oriental (north‐western Argentina) are here analysed. Richness and evenness increase through time in both deep subtidal (between fair‐weather and storm wave base) and offshore (below storm wave base) communities. Two significant patterns are superimposed on this general trend: (1) the magnitude of the increase in evenness is much more pronounced in deep than in shallower settings, and (2) richness and evenness trajectories are decoupled (while a significant rise in evenness is recorded in the middle Tremadocian (Tr2), an increase in richness is delayed until the late Tremadocian (Tr3)). In contrast to expectations, a single family (Olenidae) is dominant in samples associated with this earlier rise in evenness relative to richness. Hence, this trend is explained neither by the number of families present in the communities nor by the familial identity of the most abundant taxon. Large‐scale comparisons of the timing and geographical components of these trends are restricted to the patterns recognized in Laurentian North American studies. Results from the Cordillera Oriental mirror those of Laurentia regarding the rise in both metrics in deep marine settings. Nevertheless, the timing of this increase in richness and evenness is delayed in the Cordillera Oriental, supporting the idea that palaeogeographical regions differed in the nature and timing of ecological changes. Finally, the rise in trilobite alpha‐diversity through the Late Cambrian – Early Ordovician of the Cordillera Oriental supports the idea that trilobite alpha‐diversity did not decline worldwide, suggesting that the relative decline in trilobite alpha‐diversity is most probably caused by the dilution effect.     

Quantitatively comparing morphological trends to environment in the fossil record (Cincinnatian series; Upper Ordovician)

Determining whether morphological trends in fossil species represent evolution within a lineage or lateral shifts in morphologically variable populations through time requires a thorough examination of the details of both morphology and paleoenvironment in time and space. The purpose of this study is to explore at high resolution the relationship between morphology of the trilobite Flexicalymene granulosa and paleoenvironmental conditions in Upper Ordovician deposits of southwestern Ohio and northern Kentucky. This is achieved by using geometric morphometrics to measure high-resolution morphological changes and by using gradient analysis to capture environmental gradients underlying faunal distribution patterns. Quantitatively comparing the outcomes of these two techniques provides an assessment of whether shape changes relates to environment. Results indicate that a significant amount of shape change, seen as an anteromedial movement of the eye region over time, corresponds to ordination scores. This suggests a relationship between certain aspects of morphology and environment. The combination of these quantitative techniques has provided the foundation for determining whether morphological trends within F. granulosa represent evolution or temporal shifts in geographic variation. Future work will involve examining this relationship in greater detail both geographically and stratigraphically.     

Biogeochemical inferences of mobility of early Holocene fisher‐foragers from the Southern Sahara Desert

North Africa is increasingly seen as an important context for understanding modern human evolution and reconstructing biocultural adaptations. The Sahara, in particular, witnessed a fluorescence of hunter‐gatherer settlement at the onset of the Holocene after an extended occupational hiatus. Subsequent subsistence changes through the Holocene are contrary to those documented in other areas where mobile foraging gave way to settled agricultural village life. In North Africa, extractive fishing and hunting was supplanted by cattle and caprine pastoralism under deteriorating climatic conditions. Therefore, the initial stage of food production in North Africa witnessed a likely increase in mobility. However, there are few studies of paleomobility in Early Holocene hunter‐gatherer Saharan populations and the degree of mobility is generally assumed. Here, we present radiogenic strontium isotope ratios from Early Holocene fisher‐forager peoples from the site of Gobero, central Niger, southern Sahara Desert. Data indicate a relatively homogeneous radiogenic strontium isotope signature for this hunter‐gather population with limited variability exhibited throughout the life course or among different individuals. Although the overall signature was local, some variation in the radiogenic strontium isotope data likely reflects transhumance into the nearby Aïr Massif. Data from Gobero were significantly less variable than in other worldwide hunter‐gatherer populations, including those thought to be fairly sedentary. Strontium data from Gobero were also significantly different from contemporaneous sites in southwestern Libya. These patterns are discussed with respect to archaeological models of community organization and technological evolution. Am J Phys Anthropol 2011. © 2011 Wiley‐Liss, Inc.     

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.     

The palaeobiology of belemnites – foundation for the interpretation of rostrum geochemistry

Belemnites are an extinct group of Mesozoic coleoid cephalopods with a fossil record ranging from the early Late Triassic [about 240 million years ago (Mya)] to the Cretaceous/Palaeogene boundary (65 Mya). Belemnites were widely distributed, highly abundant and diverse, and an important component of Mesozoic marine food webs. Their internal shells, specifically their low‐Mg calcite rostra, have been used as palaeoenvironmental carbonate archives for the last 70 years. This is primarily due to the assumption that the rostrum calcite formed in equilibrium with the oxygen isotope composition of ambient sea water. Of prime importance for the reliable interpretation of isotope data derived from these biogenic carbonates is a robust reconstruction of the palaeobiology of their producers. Here we provide a critical assessment of published reconstructions of belemnite soft‐body organization and their lifestyle and habitats. Different lines of evidence, including sedimentological, geochemical, morphological, and biomechanical data, point towards an outer shelf habitat of belemnites, for some taxa also including the littoral area. Belemnite habitat temperatures, oxygen content, salinities, and life span are constrained based on observations of the ecology and life history of modern coleoids. Belemnite habitat depth might have been largely controlled by food and temperature, with a temperature optimum between 10°C and 30°C. The distribution of modern coleoids is for most species restricted to well‐oxygenated water masses and a salinity between 27 and 37 psu. The trophic position of belemnites as both predators and prey is documented by unique fossil finds of stomach contents and soft tissue preservation, such as jaws, hooks, and ink sacs. Belemnites were medium‐sized predators in the epipelagic zone (not deeper than ～200 m) hunting for crustaceans, other cephalopods, and fishes. Taxa with elongated rostra probably were fast and highly manoeuvrable swimmers. Forms with conical rostra represent slow but highly manoeuvrable swimmers, and forms with depressed rostra likely had a bottom‐related life habit. Predators of adult belemnites were sharks, bony fishes, and marine reptiles. Belemnites, like most of the modern coleoids, were relatively short lived, most likely living only for 1–2 years. Understanding the biomineralization of belemnite rostra is highly relevant for an improved interpretation of their geochemistry. Here we confirm that belemnite rostra are composed of low Mg‐calcite fibres, but they do not contain distinct types of laminae. These fibres are composed of two distinct calcite phases. One phase is a filigree network of tetrahedral organic‐rich calcite and the second phase is represented by organic‐poor calcite.     

Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management

Plant species richness and range-size rarity in Africa south of the Sahara is concentrated in centres of plant diversity and endemism. Distribution patterns of plants mapped in the Distributiones Plantarum Africanum series and selected taxonomic monographs are analysed using the computer programme WORLDMAP. The plants are divided into four groups: herbaceous geophytes, mesophytic herbs, light-demanding shrubs and woody genera. Each group has peaks of species richness and range-size rarity at locations different to the other groups. Herbaceous geophytes and mesophytic herbs have their peaks of species richness and range-size rarity in the same location, the western Cape for geophytes and the Crystal Mountain for mesophytic herbs, whereas light-demanding shrubs and woody genera have peaks in different places. The results are discussed in relation to possible factors determining species richness and endemism and their likely conservation significance.     

The graptolite correlation of the North American Upper Ordovician Standard

The North American Upper Ordovician reference standard, the Cincinnatian Series, contains rich shelly microfossil faunas in its type area in the Cincinnati Region but graptolites are uncommon in most of its shallow-water calcareous sediments. Consequently, the graptolite correlation of this key sequence has remained uncertain, in part, even controversial. A review of both previously published recently discovered graptolite Occurrences in the type Cincinnatian, combined with data from the important graptolite successions in Oklahoma New York-Quebec, has not only clarified the graptolite correlation of the Cincinnatian but also added new data on the morphology taxonomy, the vertical horizontal distribution, of several taxa. The information now at hand indicates that the Edenian Stage correlates with the C. spiniferus Zone, the Maysvillian Stage with the C. pygmaeus lower middle P. manitoulinensis Zone, the Richmondian Stage with the upper P. manitoulinensis , the D. complanatus , possibly part of the C. inuiti Zone. Comparison between graptolite conodont biostratigraphic evidence reveals no apparent conflict. Correlations are proposed between Upper Ordovician North American stages, graptolite conodont zones, successions in Texas, Oklahoma, Sweden, European graptolite zones, British series.     

Comparative sequence stratigraphy of two classic Upper Ordovician successions, Trenton Shelf (New York–Ontario) and Lexington Platform (Kentucky–Ohio): implications for eustasy and local tectonism in eastern Laurentia

Comparison of the classic Upper Ordovician (Mohawkian to lower Cincinnatian; Caradoc to lower Ashgill) Black River and Trenton Groups in New York State/southern Ontario with the Tyrone–Lexington Formations exposed in the Jessamine Dome (southern Cincinnati Arch) in north-central Kentucky and southern Ohio reveals striking similarities. Previous emphasis on complex local facies mosaic has obscured widespread regional patterns. Biostratigraphy and K-bentonites provide broad constraints on inter-regional correlations; however, an allostratigraphic approach permits higher resolution correlations and a partial test of eustatic vs. strictly local tectonic models to explain stratigraphic patterns. Upper Mohawkian to lower Cincinnatian (455–449 Ma) depositional sequences, previously recognized for the Jessamine Dome and Nashville Dome areas, are correlated between the two main study areas and further refined; Chatfieldian (Rocklandian to Shermanian or Cobourgian of traditional terminology) sequences M5 and M6 of previous workers are interpreted to be composite sequences and are each subdivided into three smaller-scale sequences, which also have counterparts in the New York–Ontario strata. In turn, these correlations indicate at least partial allocyclic control on sedimentary cycles. Complex lateral variations within depositional sequences, especially in the late Shermanian to Edenian, indicate that tectonically controlled patterns of basinal subsidence and uplift of crustal blocks (perhaps reflecting forebulge migration) exerted a strong influence on the local facies and motif of depositional sequences. These tectonic features, however, did not obliterate the underlying allocyclic pattern. Indeed, high-resolution sequence stratigraphy enables detailed resolution of shifting patterns of minor uplift and subsidence.     

Changing patterns of mobility as a response to climatic deterioration and aridification in the middle Holocene southern Sahara

The African Humid Period witnessed a rapid human re‐occupation of the Sahara as numerous lakes formed during the Holocene climatic optimum circa 10‐5 kya. Permanent waters attracted a variety of aquatic and terrestrial fauna allowing for long‐term occupation of specific paleolake basins. The Gobero paleolake in central Niger was one such location that preserves a unique mortuary record from the southern Sahara. Here, we use radiogenic strontium isotope analysis to investigate how human communities adapted to aridification throughout the Holocene. In particular, we examine the effects of increasing climate instability on patterns of human mobility. Results of radiogenic strontium isotope analysis of enamel and bone samples from Middle Holocene burials (～7.2–4.9 kya) indicate predominantly local values with no evidence for sex‐based variation. Comparisons of radiogenic strontium isotope data with previously published (Stojanowski and Knudson: Am J Phys Anthropol 146 (2011) 49–61) Early Holocene burials (～9.7–8.3 kya) indicate significant differences in both enamel and bone values. Middle Holocene individuals demonstrate a predominantly non‐local signature for enamel values and a predominantly local signature for bone values. Those individuals with non‐local bone values always demonstrated non‐local enamel values; however, the opposite was not the case. This suggests a divergence of mobility strategies during the Middle Holocene with a minority of individuals maintaining a more mobile existence throughout their life and others maintaining a similar strategy as Early Holocene hunter–gatherers that was tied to the paleolake basin. The more mobile individuals likely lived during the terminal phase of the lake's occupation. One response to aridification by Saharan peoples, then, was increasing mobility. Am J Phys Anthropol 154:79–93, 2014. © 2014 Wiley Periodicals, Inc.     

Evolution of the Rhynchotrema–Hiscobeccus lineage: implications for the diversification of the Late Ordovician epicontinental brachiopod fauna of Laurentia

Multivariate analysis based on nine biometric characters of 171 Late Ordovician rhynchonellide specimens from nine upper Sandbian–upper Katian localities in North America supports the hypothesis that one of the diagnostic taxa of the North American epicontinental brachiopod fauna, Hiscobeccus, evolved from Rhynchotrema, which lived predominantly in peri‐cratonic settings. The oldest known Hiscobeccus, H. mackenziensis of early Katian age, exhibits transitional characteristics between Rhynchotrema and Hiscobeccus, and it clusters more closely with Rhynchotrema than with younger species of Hiscobeccus of mid–late Katian (Maysvillian–Richmondian) age. Diversification of the Hiscobeccus lineage in epicontinental seas was characterized by drastic increase in shell size, globosity and lamellosity, especially in palaeoequatorially located inland basins. Such morphological trends are interpreted as adaptation to relatively shallow, muddy substrates, moderate water turbulence, relatively low oxygen content and unstable supply of nutrients in generally overheated epicontinental seas with sluggish circulation.