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Front Matter
Dedication to Calvin H. Stevens
Dedication to James H. Trexler Jr.
Late Paleozoic and Early Mesozoic Tectonostratigraphy and Biostratigraphy Of Western Pangea—Volume Overview
Late Paleozoic Tectonostratigraphic Framework of the Western North America Continental Margin
The late Paleozoic continental margin of western Pangea was in tectonic flux from at least the mid-Paleozoic Antler orogeny through the Late Permian–earliest Triassic Sonoma orogeny. This tectonism is registered by the periodic and apparent synchronous initiation and/or disruption of sedimentary basins and their associated paleogeographic highs along the entire length of the continental margin. The continental margin was not “passive” during the late Paleozoic, as is commonly believed. The possible tectonic drive(s) for this tectonism are problematic and include (1) terrane–continent collisions, (2) transpression and transtension along a long-lived translational margin, (3) far-field stresses related to continent–continent collision along the Appalachian–Ouachita–Marathon margins, and (4) shifts in mantle-plate interactions and resulting changes in global plate motions and intraplate stresses. Regardless of the specific tectonic driver, it must encompass the periodic and synchronous nature of these tectonic events and factor in the influence of preexisting crustal structures.
The Gzhelian (Upper Pennsylvanian) to Kungurian (Lower Permian) succession around Carlin Canyon, northern Nevada, in the Basin and Range province of the western USA is a relatively undeformed wedge of fossiliferous marine carbonate and fine-grained calcareous and cherty clastic rocks that rests with profound angular unconformity on Mississippian to mid-Pennsylvanian sedimentary rocks that had been uplifted, faulted, folded, and eroded prior to the Late Pennsylvanian transgression. This wedge of sediments, which tapers over less than 2 km from 1341 m in the west to 588 m in the east, comprises the Strathearn, Buckskin Mountain, and lower part of the Beacon Flat formations. These units form a second-order sequence within which five third-order unconformity-bounded transgressive–regressive sequences are nested. These sequences are Gzhelian, early to late Asselian, latest Asselian to late Sakmarian, latest Sakmarian to late Artinskian, and latest Artinskian to late Kungurian in age based on the determination and biostratigraphic interpretation of 26 conodont taxa, including two new species ( Adetognathus carlinensis n. sp. and Sweetognathus trexleri n. sp.). Each sequence records sedimentation on a westward-dipping ramp along which significant facies change occurs with inner-ramp coarse-grained algal and bioclastic photozoan grainstone to the east passing westward into mid- to outer-ramp heterozoan carbonate, and ultimately into deep-water fine-grained mixed clastic–carbonate facies with no fossils except sponge spicules, representing deep-water sedimentation in a basinal area that underwent repeated episodes of rapid subsidence associated with each sequence. Accommodation during sedimentation of Gzhelian–Kungurian sequences around Carlin Canyon was repeatedly created in response to flexural subsidence caused by tectonic loading west of the study area. Each sequence recorded the simultaneous foundering of the basinal area in the west and uplift of the basin margin in the east. Individual sequences overlap the underlying sequence to the east, while flexural subsidence from the Gzhelian to the earliest Artinskian led to a basin in the west that became deeper over time. A lull in tectonic activity associated with each sequence allowed carbonates to prograde from east to west, partially filling the basinal area until the early Artinskian, and completely filling it to sea level during the late Artinskian and then again in the late Kungurian. The Gzhelian–Kungurian carbonate succession of the Carlin Canyon area bears much resemblance with the coeval succession that occurs all along the northwest margin of Pangea, from Nevada in the south to the Canadian Arctic islands in the north, and down from the Barents Sea to the central Urals to the east. That broad area was affected by the same oceanographic events, the most significant of which was the earliest Sakmarian closure of the Uralian seaway, which prevented warm water from the Tethys Ocean from reaching the northwestern Pangea margin as it did before; this led to much cooler oceanic conditions all along western North America, even in the low tropical paleolatitudes where northern Nevada was located, in spite of a globally warming climate following the end of the late Paleozoic ice age.
Mississippian sedimentary facies belts in east-central California, occurring primarily in the autochthon (lower plate) of the Last Chance Thrust, are consistently oriented in a northeast–southwest direction. The boundary of one belt is marked by the depositional limit of the Osagean to Meramecian Santa Rosa Hills Limestone; a second belt farther to the northwest is bordered by the erosional truncation of the Kinderhookian to Osagean Tin Mountain Limestone. Two additional facies belts, both in the Meramecian to Chesterian Kearsarge Formation, also are present in the area; one near Jackass Flats is marked by the presence of limestone and quartzite olistoliths, and the other in the Last Chance Range includes abundant chert–pebble conglomerates. These two facies of the Kearsarge Formation also occur to the southwest at and near Mazourka Canyon in the allochthon (upper plate) of the Last Chance Thrust. The great similarity and near alignment of these facies belts in both the allochthon and the autochthon can be explained by clockwise rotation of ~55° of the allochthon around a pivot point in the west-central Inyo Mountains. In this model, displacement on the Last Chance Thrust increases from zero at the pivot point to 75 km for rocks exposed in the northern White Mountains. Reconstruction of the paleogeography suggests that the Last Chance Thrust is not part of a major fold and thrust belt but is a major structure limited to a relatively small area along the continental margin where the leading edge of an allochthonous terrane (possibly the Northern Sierra Terrane) impinged against the North American plate.
The concept of the Permian Last Chance Thrust has passed through many stages. Here we point out which critical observations have led to changes in the way this important feature has been interpreted.
Detailed mapping and reevaluation of biostratigraphic data provide new insights into the regional stratigraphic significance of the Ordovician Comus Formation at its type locality at Iron Point, Edna Mountain, Humboldt County, Nevada. Mapping of the internal stratigraphy of the Comus Formation yielded six new subunits and a previously unrecognized formation that is potentially correlative to the Middle Ordovician Eureka Quartzite. The age designation of the Comus Formation was reexamined, using the most current understanding of Ordovician graptolite biostratigraphy. The species of graptolites found in the Comus strata at Iron Point are Late Ordovician, in contrast to the Middle Ordovician age assignment in previous studies. Structural analyses using the new detailed mapping revealed six deformational events at Iron Point. The first fold set, F 1 , is west-vergent and likely correlative to mid-Pennsylvanian folds observed nearby at Edna Mountain. The second fold set, F 2 , records north–south contraction and is likely correlative to Early Permian folds observed at Edna Mountain. The King fault is a normal fault that strikes north and dips east. It truncates the F 1 and F 2 fold sets and has not been active since the Early Permian. The Silver Coin thrust strikes east, places the Ordovician Vinini Formation over the Comus Formation, truncates the King fault, and is not affected by the F 1 and F 2 fold sets. Timing of the Silver Coin thrust is unknown, but it is likely post-Early Permian based on crosscutting relationships. The West fault strikes southeast and dips southwest. It truncates the Silver Coin thrust on the west, and the fault surface records several phases of motion. Finally, Iron Point is bounded on the east side by the Pumpernickel fault, a normal fault that strikes north and dips east. The movement on this structure is likely related to Miocene to Recent Basin and Range faulting. Several key findings resulted from this detailed study of the Ordovician rocks at Iron Point. (1) Based on detailed mapping of the internal stratigraphy of the Comus Formation at Iron Point, it is here interpreted to be correlative with the autochthonous Late Ordovician Hanson Creek Formation rather than the well-known “Comus Formation” that hosts Carlin-style gold mineralization in the Osgood Mountains to the north. (2) The Comus Formation at Iron Point is autochthonous, and the Roberts Mountains thrust is not present at Iron Point, either at the surface or in the subsurface. (3) The stratigraphic mismatch between Iron Point and Edna Mountain requires a fault with significant lateral offset between the two areas; its current expression could be the West fault. (4) West- and southwest-vergent structures at Iron Point and Edna Mountain are rotated counterclockwise relative to northwest-vergent structures at Carlin Canyon and elsewhere in northern Nevada. This relationship is consistent with large-scale sinistral slip along the continental margin to the west.
Structures at Buck Mountain, Nevada: Establishing the Southeastern Extent of Mid-Pennsylvanian Tectonism
This paper reports the structural and stratigraphic history of Buck Mountain, Nevada, and its regional significance in the development of southwestern Laurentia during the late Paleozoic. The two distinct generations of folding have similar style and/or timing to other fold sets in late Paleozoic strata of northern Nevada. Unconformities in the upper Paleozoic strata at Buck Mountain are consistent with unconformities documented in northern and east-central Nevada. Northwest-vergent folds (F 1 ) in the Morrowan–Atokan Ely Limestone are erosionally truncated and unconformably overlain by the middle Desmoinesian Hogan Formation and middle Wolfcampian (Sakmarian) Upper Strathearn Formation. This upper Paleozoic stratigraphic package was subsequently refolded by the Buck Mountain Syncline and associated mesoscale folds (F 2 ). F 2 folds lack tight age control but are interpreted to be associated with the Cretaceous central Nevada thrust belt. Critically, none of these structures are localized above or below low-angle faults. The unconformity between the Ely and Hogan formations is consistent with the C5 regional unconformity. Importantly, it constrains the age of northwest-vergent deformation on Buck Mountain. West-vergent folds and west-directed thrusts are documented at several locations in northern and east-central Nevada, but because of the dominance of the C6 unconformity and/or lack of robust age control, the age of these structures has not been tightly constrained. The evidence at Buck Mountain indicates that west-vergent structures predate the C5 unconformity. Buck Mountain is important because it: (1) precisely brackets the age of west-vergent deformation in Nevada to pre–mid-Desmoinesian (sub C5-unconformity) and (2) defines a southeastern edge to the late Paleozoic west-vergent deformation in northern and east-central Nevada.
Late Paleozoic Shortening in South-Central Nevada and Regional Correlations of Major Pre-Sevier Structures
Recent tectonic reinterpretations of the Late Paleozoic Southwest Laurentian margins recognize widespread Late Paleozoic deformation as a critical component in the boundary region development. Overprinted late Paleozoic structures record repeated shortening events in both northern and southern Nevada, but spatial and temporal data are currently lacking to resolve the evolution of this margin. The Timpahute Range, south-central Nevada, bridges part of the spatial gap between previous detailed studies of Late Paleozoic deformation. The purpose here is to (1) evaluate structures in the area that do not appear to fit with recognized Sevier hinterland structures (the Central Nevada thrust belt [CNTB]) and (2) consider whether these contractional structures may be Late Paleozoic and possibly link, or not, structures to the north and south. New mapping in the Timpahute Range documents four geometrically or kinematically distinct sets of structures: Tempiute Ridge folds, Schofield Pass fault zone (SPFZ), structures of the CNTB, and Cenozoic extensional faults. The first three are interpreted to represent separate shortening events based on cross-cutting relations and differences in orientations of the Tempiute Ridge folds and SPFZ (north [N]), and structures of the CNTB (northwest [NW]). The Tempiute Ridge folds represent the oldest event, D 1 . These folds are large, trend N and verge east (E). The SPFZ is west (W)-vergent, cuts across the limb of a D 1 fold and represents D 2 . The SPFZ is interpreted to be older than the CNTB structures, D 3 , based on positions of fault cut offs, and differences in footwall and hangingwall facies. All of the shortening events predate the newly dated 102.9 ± 3.2 Ma Lincoln stock and its contact metamorphic aureole. New and previous correlations suggest that a belt of Permian deformation extends from southeast (SE) California northward at least to the Timpahute Range. The Tempiute Ridge folds and SPFZ have the same distinctive geometries, styles, and kinematics as structures in the Nevada National Security Site. The mountain-size, E-vergent Tempiute Ridge folds and the W-vergent SPFZ correlate to structures associated with the Belted Range thrust and the W-vergent CP thrust, respectively. The Belted Range thrust previously has been correlated southward into the Death Valley region. Thus, convergence created large-amplitude folds and thrusts for ~200 km along strike. Structures of this age are exposed in northern Nevada but are smaller. These new relations fill a data gap and suggest differences in the size and structural style of Permian structures along strike and corresponding variations in the plate boundary configuration.
Evolution of the Pennsylvanian Ely–Bird Spring Basin: Insights from Carbon Isotope Stratigraphy
Analysis and correlation of strata in ancient basins are commonly difficult due to a lack of high-resolution age control. This study tackled this problem for the latest Mississippian to middle Pennsylvanian Ely–Bird Spring basin. Here, 1095 new carbon isotope analyses combined with existing biostratigraphy at six sections throughout the basin constrain changes in relative sediment accumulation rates in time and space. The Ely–Bird Spring basin contains dominantly shallow-water carbonates exposed in eastern and southern Nevada, western Utah, and southeastern California. It formed as part of the complex late Paleozoic southwestern Laurentian plate margin. However, the detailed evolution of the basin, and hence the tectonic driver(s) of deformation, is poorly understood. The combined isotopic and biostratigraphic data were correlated using the Match-2.3 dynamic programming algorithm. The correlations show a complex picture of sediment accumulation throughout the life of the Ely–Bird Spring basin. Initially, the most rapid sediment accumulation was in the eastern part of the basin. Throughout Morrowan time, the most rapid sediment accumulation migrated to the northwestern part of the basin, culminating in a peak of sediment accumulation in Atokan time. This peak records tectonic loading at the north or northwest margin of the basin. Basin sedimentation was interrupted by early Desmoinesian time in the north by formation of northwest-directed thrust faults, folds, uplift, and an associated unconformity. Deposition continued in the south with a correlative conformity and increased clastic input. The combination of isotopic and biostratigraphic data for correlation is therefore a valuable tool for elucidating temporal basin evolution and can be readily applied to tectonically complex carbonate basins worldwide.
The stratal architecture of the upper Ely Limestone and Mormon Gap Formation (Pennsylvanian–Lower Permian) in west-central Utah reflects the interaction of icehouse sea-level change and tectonic activity in the distal Antler–Sonoma foreland basin. Nineteen stratigraphic sections correlated by physical and biostratigraphic means provide a basis for tracing Carboniferous–Permian boundary strata over a north–south distance of 60 km. These formations can be subdivided into 14 unconformity-bounded, third-order depositional sequences of differing internal architecture and regional extent. Conodonts and fusulinids provide ages for selected sequences and parasequences, permitting correlation with tectonostratigraphic units in the proximal foreland in north-central Nevada and with selected Midcontinent cyclothems. The 14 third-order sequences stack into three second-order supersequences characterized by distinctive differences in facies and facies stacking patterns, regional continuity of cycles, relative abundance of dolomite and limestone, calculated rock accumulation rates, and the frequency and inferred duration of sequence-bounding hiatuses. These reflect the effect of high-frequency sea-level change on an intermittently subsiding distal foreland shelf. Sediment accommodation was relatively high during the Bashkirian through middle Moscovian (upper part of Lower Absaroka I supersequence) and again during the late Sakmarian and Artinskian (lower part of Lower Absaroka III supersequence) as a function of continuous subsidence and high-amplitude sea-level change. During the late Moscovian through upper Sakmarian (Lower Absaroka II supersequence), however, subsidence slowed or ceased in response to tectonic activity in north-central Nevada, with concomitant development of the West-Central Utah Highlands (forebulge). During this episode of reduced subsidence, intermittent sedimentation was driven by second- and third-order eustatic fluctuations in sea level. Constituent strata form a wedge of onlapping, northward-thinning sequences and parasequences deposited during selected third-order highstands of the Lower Absaroka II second-order sea-level event. Depositional sequences in the distal foreland are bounded by low-relief disconformities of variable duration, in contrast to the angular unconformities and intensely deformed tectonostratigraphic domains that characterize the proximal foreland basin in north-central Nevada.
Sonoma Orogeny—A Reassessment
The Late Permian to earliest Triassic Sonoma orogeny has long been envisioned as the result of an arc-continent collision that closed the Havallah oceanic basin, creating the Golconda allochthon, which was emplaced eastward onto the western edge of the continental margin along the Golconda thrust. Critical reevaluation of available stratigraphic, biostratigraphic, and structural data raise some fundamental issues with this scenario, including: (1) The Golconda allochthon experienced multiple phases of deformation both older and younger than the Sonoma orogeny; (2) the tectonostratigraphic successions in the Golconda allochthon record a disrupted depositional history; (3) these punctuated events and unconformities are mirrored by simultaneous punctuated tectonic disruptions of the adjacent continental margin; (4) some of the lithotectonic units within the Golconda allochthon have clear ties to a magmatic arc. These observations indicated that the Havallah basin did not originate as a simple, post-Antler orogeny rift basin, nor is the Mediterranean model for opening of a basin a solution to the initiation of this basin. Instead they imply a more complex paleogeography for the Havallah basin. The Late Permian–earliest Triassic closure of the Havallah basin did result in the development of the Golconda allochthon sensu stricto , but final emplacement of the Golconda allochthon was likely an Early–Middle Jurassic event.
Paleogeographic Implications of Open-Marine Anoxia in the Permian–Triassic Slide Mountain Ocean
The end-Permian mass extinction was associated with the onset of anoxia in widespread marine environments; however, the extent of this anoxia remains controversial. Proposed models range from near-universal “superanoxia” in the Panthalassic Basin to a more limited expansion of anoxia in the upper water column in response to enhanced primary productivity. The Peck Creek and Ursula Creek sections of northern British Columbia were deposited at ~200 m water depth in the Ishbel Trough, on the margin of cratonic North America. This trough was generally contiguous with the Slide Mountain Ocean, and thus with the broader Panthalassic Ocean, though it may have been partially separated by structural highs at various times during the Permian. Both sections include continuous Wordian to Changhsingian sedimentary successions, which span the end-Permian mass extinction boundary and continue into the earliest Triassic. The extinction is recognizable as the disappearance of biogenic silica from the environment, which defines the contact between the Fantasque Formation and overlying Grayling Formation. This surface also corresponds with the onset of anoxia, and the accumulation of redox-sensitive trace metals. The covariation trends in these metals, and in other isotopic proxies, can be used as tools to trace the degree of communication between the Ishbel Trough–Slide Mountain Ocean and the broader Panthalassic Basin. Molybdenum-uranium covariation trends indicate that the northern Slide Mountain Ocean and Ishbel Trough remained in communication with the larger global ocean throughout this interval, suggesting inversion of the Wordian structural high to form a depositional subbasin by the Changhsingian. This is in contrast to the Opal Creek section of southern Alberta, which shows evidence for some degree of restriction, suggesting that the Slide Mountain Ocean may have maintained a north–south gradient in water chemistry. Several lines of evidence suggest that this onset of anoxia was not related to expansion of an upwelling-driven oxygen minimum zone. No clear changes in primary productivity, as recorded by organic carbon or authigenic phosphorus and barium, are observed across the extinction horizon. Changhsingian nitrogen isotope values are generally in the 2 to 3‰ range, suggesting minimal denitrification at thermocline water depths, and these values decreased in the earliest Triassic, likely in response to enhanced nitrogen fixation. This suggests that anoxia was driven by shoaling of a chemocline that developed due to stratification of the Slide Mountain Ocean, rather than western-boundary upwelling effects.
A thick succession of upper Paleozoic carbonate rocks and minor chert crops out north of the head of Otto Fiord (northwest [NW] Ellesmere Island, Nunavut) in the Canadian Arctic Archipelago. These rocks accumulated in a tectonic subbasin—the Otto Fiord Depression (OFD)—of the Sverdrup Basin that likely originated through rifting during late Early Carboniferous (Serpukhovian). Following a long interval of passive subsidence that allowed a thick succession of Moscovian–Kasimovian carbonate rocks to fill the OFD, tectonic activity resumed during the Gzhelian (Late Pennsylvanian). This resulted in rapid collapse of the depression along its axis and simultaneous uplifts of its margins, a style of tectonism in accord with the inferred basin-wide shift to a transpressional–transtensional stress regime at that time. Late Pennsylvanian–Early Permian sedimentation in the OFD led to the development of four long-term (second-order) transgressive–regressive sequences of early Gzhelian–middle Asselian (<1200 m), late Asselian–late Sakmarian (<380 m), latest Sakmarian–late Artinskian (<160 m) and latest Artinskian–late Kungurian (<60 m) age. These ages are supported by integration of biostratigraphic data from conodonts, fusulinaceans, and small foraminifers. The development of each sequence-bounding unconformity was associated with renewed tectonism in the OFD. Each sequence recorded the development of a depositional system characterized by high energy peripheral shoreface grainstones passing basinward across a gently dipping ramp into deep-water basinal calcareous and siliceous mudstone. The ramp portion of the early Gzhelian–middle Asselian system comprises both cool-heterozoan to warmphotozoan carbonates (Nansen Formation) suggesting a relatively shallow thermocline at that time. These rocks are arranged in a series of high-order cyclothems of glacio-eustatic origin. Cyclothemic sedimentation ended at the Asselian–Sakmarian boundary, simultaneous to a major depositional system shift to cool-water heterozoan sedimentation (Raanes Formation), a change presumably brought on by the closure of the Uralian seaway linking NW Pangea with the Tethyan Ocean. This event led to the destruction of the permanent thermocline, and disappearance of photozoan carbonates by the early Sakmarian despite rising temperatures globally. Cool-water heterozoan sedimentation, associated with relatively shallow outer-ramp to midramp spiculitic chert resumed in the Artinskian and then again in the Kungurian (Great Bear Cape Formation) when the OFD was filled up. The depression ceased to exist as a separate tectonic/subsidence entity with the widespread sub-Middle Permian unconformity, above which sediments were deposited during a passive subsidence regime across most of the Sverdrup Basin. The Pennsylvanian–Lower Permian succession that accumulated in the OFD along the clastic-free northern margin of the Sverdrup Basin is essentially identical, both in terms of tectonic evolution and stratigraphic development, with the coeval succession of Raanes Peninsula, southwest (SW) Ellesmere Island, the type area of the Raanes, Trappers Cove, and Great Bear Cape formations along the clastic-influenced southern margin.
This article utilizes over 3000 biostratigraphic reports of fusulinid taxa in the Midland Basin to produce a series of chronostratigraphic surfaces that show shelf-to-basin profiles from the end of the Atokan to the end of the Leonardian. The position of the shelf-edge break along the Eastern Shelf was geometrically reconstructed for each of the chronostratigraphic surfaces. Comparing the location of the shelf edges produced in this study to published examples resulted in significant disagreement for some time intervals, especially the Wolfcampian. These discrepancies are inferred to be predominantly the result of lithostratigraphic-based vs. biostratigraphic-based data. Assessing shelf-edge trajectory through the Pennsylvanian into the early Permian indicates that (1) tectonic and eustatic increases in shelf accommodation resulted in retrogradation of the shelf edge during the Pennsylvanian, and (2) early Permian progradation was the likely result of cessation in tectonic subsidence that allowed bypass of the shelf and passive filling of the basin center. When placed into the context of the Ancestral Rocky Mountains, subsidence analysis of the Midland Basin agrees with tectonic models that portray a synchronous start rather than an east-to-west migration of peak subsidence. Additionally, a relatively synchronous apex of tectonic subsidence occurred in the Middle to Late Pennsylvanian.
The Ancestral Rocky Mountains (ARM) represent an intraplate deformational event that resulted in a series of Precambrian-cored basement uplifts with adjacent basins that accumulated Pennsylvanian to early Permian strata. Tectonic models for the event are debated largely because of the lack of robust age control in the basin fill. In New Mexico, the ARM event resulted in a series of basins with some of the best biostratigraphic records across the orogenic province. This article utilizes published biostratigraphic and lithostratigraphic data to (1) reconcile the onset of subsidence by first accumulation of Pennsylvanian strata, (2) establish a period of peak subsidence estimated by maximum accumulation rate, (3) correlate estimated peak subsidence with the first appearance of arkose derived from the adjacent denuded Precambrian-cored block, and (4) demarcate synorogenic strata from Permian strata that are postorogenic. Results demonstrate that within New Mexico ARM basins, (1) onset was relatively synchronous, predominantly beginning in early Atokan time; (2) peak subsidence, while potentially younging southward, was relatively coeval in the Middle to Late Pennsylvanian; (3) first occurrence of arkose either predates the period of peak subsidence or is coeval with peak subsidence; and (4) early Permian strata across the study area onlap preexisting faults and folds, and/or form a buttress unconformity with Precambrian basement.
Carboniferous conodont biostratigraphy
Abstract Carboniferous conodont biostratigraphy comprises regional zonations that reflect the palaeogeographical distribution of taxa and distinct shallow-water and deep-water conodont biofacies. Some species have a global distribution and can effect high quality correlations. These taxa are incorporated into definitions of global Carboniferous chronostratigraphic units. A standard global Carboniferous zonation has not been developed. The lowermost Mississippian is zoned by Siphonodella species, excepet in shallow-water facies, where other polygnathids are used. Gnathodus species radiated during the Tournaisian and are used to define many Mississippian zones. A late Tournaisian maximum in diversity, characterized by short-lived genera, was followed by lower diversity faunas of Gnathodus species and carminate genera through the Visean and Serpukhovian. By the late Visean and Serpukhovian, Lochriea provides better biostratigraphic resolution. Shallow-water zonations based on Cavusgnathus and Mestognathus are difficult to correlate. An extinction event near the base of the Pennsylvanian was followed by the appearance of new gnathodid genera: Rhachistognathus , Declinognathodus , Neognathodus , Idiognathoides and Idiognathodus . By the middle of the Moscovian, few genera remained: Idiognathodus , Neognathodus and Swadelina. During the middle Kasimovian and Gzhelian, only Idiognathodus and Streptognathodus species were common. Near the end of the Gzhelian, a rediversification of Streptognathodus species extended into the Cisuralian.