John R. Groves

Foraminiferal diversification during the late Paleozoic ice age

Abstract A record of late Paleozoic foraminiferal diversity, origination and extinction frequencies, and provincialism at million-year temporal resolution and species-level taxonomic resolution has been achieved by analyzing composite standard databases. Foraminiferal species diversity increased throughout Mississippian and Pennsylvanian time leading up to its peak at the Pennsylvanian/Permian boundary. Foraminifers then experienced a steep decline in diversity during the Early Permian. Frequencies of origination and extinction broadly tracked changes in global diversity. From Late Mississippian time onward, patterns in total foraminiferal diversity were dominated by fusulinoideans. There is no clear relationship between rates of foraminiferal evolution and the alternating glacial and nonglacial intervals that characterized the late Paleozoic ice age. Rather, high rates of origination and extinction might reflect instability of neritic environments as a consequence of high-frequency, high-amplitude base-level fluctuations (cyclothemic deposition). Further, the advent of algal symbiosis in fusulinoideans was a physiologic innovation that promoted diversification as these symbiont-bearing taxa experimented with morphologic adaptations for partitioning the low-nutrient environments to which they were specialized. Growth to large size and delayed maturation in fusulinoideans might have been enabled by the late Paleozoic hyperoxic atmosphere and the widespread development of oligotrophic, carbonate platform and shelf environments. The late Paleozoic history of foraminiferal diversification was influenced also by closure of the Rheic Ocean beginning in Late Mississippian time. Foraminiferal associations on opposite sides of Pangea exhibited relatively high similarity prior to the closure, but then similarity decreased steadily after destruction of the subequatorial marine corridor. Arctic-Eurasian and North American associations were nearly isolated from one another throughout the main burst of fusulinoidean diversification, so that parallel lineages developed independently in the two regions, resulting in many instances of convergence.

Late paleozoic fusulinoidean gigantism driven by atmospheric hyperoxia.

Atmospheric hyperoxia, with pO2 in excess of 30%, has long been hypothesized to account for late Paleozoic (360–250 million years ago) gigantism in numerous higher taxa. However, this hypothesis has not been evaluated statistically because comprehensive size data have not been compiled previously at sufficient temporal resolution to permit quantitative analysis. In this study, we test the hyperoxia‐gigantism hypothesis by examining the fossil record of fusulinoidean foraminifers, a dramatic example of protistan gigantism with some individuals exceeding 10 cm in length and exceeding their relatives by six orders of magnitude in biovolume. We assembled and examined comprehensive regional and global, species‐level datasets containing 270 and 1823 species, respectively. A statistical model of size evolution forced by atmospheric pO2 is conclusively favored over alternative models based on random walks or a constant tendency toward size increase. Moreover, the ratios of volume to surface area in the largest fusulinoideans are consistent in magnitude and trend with a mathematical model based on oxygen transport limitation. We further validate the hyperoxia‐gigantism model through an examination of modern foraminiferal species living along a measured gradient in oxygen concentration. These findings provide the first quantitative confirmation of a direct connection between Paleozoic gigantism and atmospheric hyperoxia.

Timing and Size Selectivity of the Guadalupian (Middle Permian) Fusulinoidean Extinction

A comprehensive, high resolution stratigraphic database of fusulinoidean foraminifers reveals that this group of protists suffered extreme losses during the Guadalupian extinction. Most species (88%) were eliminated gradually over the course of 9 myr during the Wordian and Capitanian ages. A pulse of greatly elevated per capita extinction frequency occurred during the last million years of the Capitanian (260–259 Ma). Contrary to prevailing opinion, the end-Capitanian event did not preferentially eliminate large, morphologically complex species in the families Schwagerinidae and Neoschwagerinidae, because most species in those families were already extinct. Rather, 69 percent of the species eliminated at the end of the Capitanian were small, morphologically conservative representatives of the Ozawainellidae, Schubertellidae and Staffellidae. Survivors from these families comprised the low-diversity association of Wuchiapingian fusulinoideans. Schubertellids, and to a lesser extent ozawainellids, diversified in the late Wuchiapingian and Changhsingian ages before the final demise of fusulinoideans during the end-Permian mass extinction. The Wordian–Capitanian fusulinoidean attrition might have been caused by photosymbiont loss and habitat reduction stemming from an interval of global cooling termed the Kamura event (∼265–259.5 Ma), although the onset of fusulinoidean diversity decline predates geochemical evidence for the beginning of the Kamura event by ∼3 myr. The end-Capitanian extinction pulse might reflect environmental deterioration from the combined effects of global cooling, Emeishan effusive volcanism and sea-level lowstand.

Biostratigraphy and evolution of Late Carboniferous and Early Permian smaller foraminifers from the Barents Sea (offshore Arctic Norway)

Nearly continuous cores from a 500-meter interval of upper Moscovian through post-Artinskian carbonate strata on the Finnmark Platform have yielded rich assemblages of fusulinaceans and smaller foraminifers. The fusulinaceans provide an independent time framework for evaluating stratigraphic occurrences of associated smaller foraminifers. Information derived from this study has been integrated with that from previous investigations to produce a smaller foraminiferal biostratigraphic model for the High Arctic. Kasimovian strata are characterized by occurrences of Nodosinelloides spp., Protonodosaria spp. and Hemigordius schlumbergeri. Overlying lower Gzhelian beds are identified by the appearances of Raphconilia modificata and Amphoratheca iniqua. Tezaquina clivuli and Cribrogenerina gigas first occur in upper Gzhelian strata, and Asselian rocks contain appearances of Geinitzina postcarbonica and Pachyphloia spp. Phylogenetic reconstructions suggest that the Late Carboniferous-Early Permian pseudovidalinids (Archaediscacea) derived from the Pseudoammidiscidae, as did the older group of predominantly Early Carboniferous archaediscaceans. The repeated development of similar morphologies within the two groups of archaediscaceans includes examples of both convergence and iterative evolution. The Protonodosariidae and Syzraniidae most likely evolved from an earlandiid ancestor. The syzraniids gave rise to the Geinitzinidae, which in turn gave rise to the Pachyphloiidae. As in the archaediscaceans, evolution within and among the Earlandiidae, Protonodosariidae, Syzraniidae, Geinitzinidae and Pachyphloiidae includes multiple examples of repeated patterns. Two key phenotypic developments seemingly led to bursts of diversification in different groups throughout late Paleozoic foraminiferal evolution. The acquisition of an outer hyaline or pseudofibrous wall layer was a morphologic breakthrough followed by taxonomic radiation in both groups of archaediscaceans and in the Earlandia-Syzrania lineage. The second major event was the shift from an undivided tubular morphology to uniserial morphology in both the Earlandiidae-Protonodosariidae and Syzraniidae-Geinitzinidae lines. Nodosinelloides pinardae is proposed as a new name for Nodosaria grandis Lipina, 1949 (preoccupied).

Wall structures in selected Paleozoic Lagenide foraminifera

Abstract Paleozoic lagenide foraminifera are strikingly similar morphologically to Mesozoic and Cenozoic Lagenida, but because benthic foraminifers suffered a catastrophic reduction in diversity during the end-Permian mass extinction, it is unclear whether the similarities of Paleozoic lagenides and younger unquestioned Lagenida link them evolutionarily or are the product of convergence. Seven species representing five families of Paleozoic lagenide foraminifers were examined and found to possess mostly monolamellar hyaline-radial walls, as in extant nodosariid Lagenida. Exceptions are Protonodosaria rauserae Gerke, 1959, whose wall is not optically radial, and Syzrania amazonica Altiner and Savini, 1997, whose hyaline-radial wall may or may not be accompanied by a secreted inner microgranular layer. The inner microgranular layer is an element that is thought to have been inherited from the ancestral Fusulinida. Its absence in all but the earliest and morphologically simplest Paleozoic lagenides indicates more advanced wall structure than generally has been ascribed to this group. The wall in Pachyphloia spp. is secondarily thickened, suggesting plesio- or ortho-monolamellar construction, whereas other examined species exhibit atelo-monolamellar wall structure. These types of lamellarity are common among modern nodosariids. Thus, on the basis of not only morphologic similarity but also similar wall structure, we strongly suspect evolutionary continuity of lagenides across the Permian-Triassic boundary. The question is not completely resolved, however, because lagenides have not yet been recovered from lowest Triassic rocks.

ORIGIN AND EARLY EVOLUTIONARY RADIATION OF THE ORDER LAGENIDA (FORAMINIFERA)

Abstract The Order Lagenida is a monophyletic group of calcareous foraminifers that originated in Middle Pennsylvanian time via acquisition of hyaline-radial wall structure and loss of microgranular wall structure, the latter being characteristic of the close sister group and likely ancestor, the Fusulinida. Early lagenides are delineated into subgroups on the basis of presence or absence of partitioning within their tests, and among partitioned forms, on continuous versus discontinuous growth styles. Partitioned, discontinuously growing forms may be further delineated on the basis of test symmetry and on modifications to chamber shape and apertural complexity. Early lagenides underwent rapid taxonomic differentiation during late Moscovian and early Kasimovian time. Taxonomic differentiation was accompanied by rapid dispersal from the presumed center of origin in the midcontinent-Andean area to tropical and subtropical shelves worldwide. By Early Permian time certain lagenides were adapted to cool water paleoenvironments, as evidenced by their occurrences in high paleolatitudes and even in glaciomarine basins. Early Permian lagenides do not exhibit marked provincialism, but there is evidence for paleolatitudinal control on assemblages. The midcontinent-Andean and present Arctic areas contain similar, diverse faunas from low- to mid paleolatitudes along the western margin of Pangaea. These faunas share many elements in common with faunas from the tropical and subtropical eastern margin of Pangaea (Paleotethys). In contrast, the Europe-Urals, Siberian and Australian areas are characterized by a slightly different faunal association from mid- to high paleolatitudes in both hemispheres. Panthalssan faunas are less well known, but seemingly contain only cosmopolitan taxa.

Foraminiferal Biostratigraphy of the Visean–Serpukhovian (Mississippian) Boundary Interval At Slope and Platform Sections In Southern Guizhou (South China)

Abstract The Visean–Serpukhovian boundary is not yet defined by a Global Stratotype Section and Point (GSSP) but it is recognizable operationally by the appearance of the conodont Lochriea ziegleri in the L. nodosa–L. ziegleri chronocline. Foraminiferal successions across this boundary in the type area of the Serpukhovian Stage (Moscow Basin, Russia), elsewhere in Russia and in the central United States suggest that the appearances of Asteroarchaediscus postrugosus, Janischewskina delicata, Eolasiodiscus donbassicus, and specimens controversially referred to “Millerella tortula” are reliable, auxiliary indices to the base of the Serpukhovian. In southern Guizhou Province, China, Visean–Serpukhovian rock sequences from slope and platform settings have yielded rich associations of conodonts and foraminifers, respectively. The Nashui section is a leading candidate for the Serpukhovian GSSP because its slope deposits contain an uninterrupted record of conodont occurrences including the L. nodosa–L. ziegleri transition. Foraminifers recovered from the Nashui section are comparatively rare and include none of the basal Serpukhovian indices. In contrast, the nearby Yashui section represents a platform interior setting in which foraminifers flourished and conodonts were nearly absent. The base of the Serpukhovian at Yashui is marked approximately by the appearance of “tortula-like” specimens. Although it is not possible to correlate biostratigraphically between the Nashui and Yashui sections, the occurrence of “tortula-like” specimens at the Yashui section allows correlation with the mid-Venevian Substage of the Moscow Basin at a level coinciding with the appearance of L. ziegleri. Together, the slope and platform sections comprise an informative biostratigraphic reference area for micropaleontologic characterization of the Visean–Serpukhovian boundary in southern Guizhou.

Mississippian (Lower Carboniferous) microfossils from the Chios Mélange (Chios Island, Greece)

Abstract The Chios Melange is a thick Paleozoic wildflysch sequence that crops out on the Greek island Chios. It is composed of chert, limestone and volcanic blocks floating in a siliciclastic turbiditic matrix. New data suggest that the youngest blocks within the Chios Melange are clasts of a breccia from the Kouramia–Nenitouria area that contain conodonts of late Visean or early Serpukovian age. The conodont fauna from the breccias is characterized by the genera Gnathodus and Lochriea, which favored deeper-water, open-marine facies. Elsewhere on the island, in the Papalia-Nagos area, calcareous microfossils of middle to late Visean age have been recovered from lime grainstone beds within the Chios Melange. Allochems making up the grainstones are interpreted to have been transported into deeper-water turbiditic facies, but there is no evidence of stratigraphic reworking. Age-diagnostic calcareous microfossils include the alga Koninckopora inflata and foraminifers in the genus Paraarchaediscus. In situ Mississippian microfossils indicate that the Chios Melange is older than Pennsylvanian (Upper Carboniferous) – Permian, as previously thought. The revised age suggests that the origin of the Chios Melange may be related to the development of an accretionary prism during the Hercynian Orogeny.

Calcareous algae and associated microfossils from Mid-Carboniferous rocks in east-central Idaho

Mid-Carboniferous (upper Chesterian–lower Atokan) rocks exposed north of the Snake River Plain in east-central Idaho are assigned to the Arco Hills, Bluebird Mountain and lower Snaky Canyon formations (ascending order). In the southern Lemhi Range, calcareous algae and associated microproblematica from these rocks include representatives of at least 13 genera and genus-level taxa within the Dasycladaceae, Aoujgaliaceae, Ungdarellaceae, and incertae familiae. Local appearances of Masloviporidium delicata and Donezella lutugini are early Morrowan or younger as determined independently by studies of associated foraminifers and conodonts. Beresella polyramosa and Komia abundans are locally restricted to Atokan rocks. These findings are consistent with compiled data on worldwide stratigraphic distributions of these taxa, and suggest that certain Upper Paleozoic calcareous algae may be of limited value in biostratigraphic correlation.

Correlation of the Type Bashkirian Stage (Middle Carboniferous, South Urals) with the Morrowan and Atokan series of the midcontinental and western United States

The graphic correlation technique has been used to directly relate the stratigraphic appearances of key species in the Bashkirian Stage stratotype to those in a North American composite section. The type Bashkirian is separated from the underlying Serpukhovian Stage by an erosional unconformity and associated lacuna of undetermined, but probably minor duration. Accordingly, the base of the type Bashkirian (base of Bogdanovkian Horizon) is only slightly younger than the international mid-Carboniferous boundary. A level within the upper part of the Tashastian Horizon (Upper Bashkirian Substage) most likely correlates with the Morrowan-Atokan boundary. This level roughly coincides with a sequence boundary at the Bashkirian stratotype and with a regional unconformity in the North American midcontinent. The top of the Bashkirian Stage (top of Asatauian Horizon) is lower Atokan in North American terms. On the basis of recent 40 Ar/ 39 Ar and SHRIMP zircon geochronology studies, the age of the mid-Carboniferous boundary is estimated at 314 Ma and a horizon of early Atokan age is dated at 310.8 Ma. Accepting the present biostratigraphic correlations, these values suggest a duration for the Bashkirian Stage of slightly more than 3.2 m.y. and a duration of the Morrowan Series of slightly less than 3.2 m.y.