Biodiversity and biogeography of middle–late liassic ammonoids: implications for the early Toarcian mass extinction

In Western Tethyan areas, the Toarcian stage begins with two important evolutionary events in ammonite faunas: (1) the disruption of Tethyan–Boreal provinciality; (2) a biological crisis linked with the oceanic anoxic event OAE. The analysis of these events has been addressed by constructing curves of ammonoid diversity (species richness, origination and extinction rates) in the Late Pliensbachian (= Domerian)–Early Toarcian interval in selected localities. Two diversity drops are recognized. The first one is recorded at the end of the Dactylioceras mirabile subzone and reflects the disruption of Tethyan–Boreal provinciality, through the progressive extinction of the Boreal endemic family Amaltheidae that occupied the north-western European seas during the whole Pliensbachian on the one hand, and the extinction of Late Domerian Ammonitina endemic to the Mediterranean areas on the other hand. The Early Toarcian homogeneization of Mediterranean and north-western European ammonoid faunas was reached via elimination of both Boreal and Mediterranean endemics with differential rates of extinction in the two palaeogeographic domains and the subsequent geographical expansion of Tethyan-derived ammonoids. The second, dramatic drop in ammonite diversity in the upper part of the Dactylioceras semicelatum subzone coincided with the onset of OAE. It also affected epioceanic ammonoid clades like Phyllocerataceae and Lytocerataceae. These two drops are interpreted as two distinct extinctions and not as episodes of a single, stepwise event. Complex relations between ammonoid diversity and sea-level changes are suggested by trends in endemism, which may be reversed during either a single transgression or a single regression.     

Key innovations in Mesozoic ammonoids: the multicuspidate radula and the calcified aptychus

A nearly complete radula with seven elements per row preserved inside of an isolated, bivalved, calcitic lower jaw (= aptychus) of the Late Jurassic ammonite Aspidoceras is described from the Fossillagerstätte Painten (Bavaria, southern Germany). It is the largest known ammonite radula and the first record for the Perisphinctoidea. The multicuspidate tooth elements (ctenodont type of radula) present short cusps. Owing to significant morphological differences between known aptychophoran ammonoid radulae, their possible function is discussed, partly in comparison with modern cephalopod and gastropod radulae. Analogies between the evolution of the pharyngeal jaws of cichlid fishes and the ammonoid buccal apparatus raise the possibility that the evolution of a multicuspidate radula allowed for a functional decoupling of the aptychophoran ammonoid jaw. The radula, therefore, represents a key innovation which allowed for the evolution of the calcified lower jaws in Jurassic and Cretaceous aptychophoran ammonites. Possible triggers for this morphological change during the early Toarcian are discussed. Finally, we hypothesize potential adaptations of ammonoids to different feeding niches based on radular tooth morphologies.     

Zur paläobathymetrischen zuverlässigkeit von ammonoideen-faunen-spektren

The Jurassic ammonoid faunal spectrum of the Mediterranean faunal provinces always exhibits a large quota of Phylloceratina and Lytoceratina. As a rule, these ammonites are supposed to be index fossils of deeper-water deposits. An example from Spain suggests that this is not always the case. Nevertheless general zoogeographical aspects of ammonoid distribution, irrespective of the depth of water, still merit our attention.     

The earliest known Salamanders (Amphibia,Caudata):A record from the Middle Jurassic of England

Marmorerpeton gen. nov. represented by M. kermackisp. nov. and M. freemani sp. nov., is the earliest known genus of fossil salamander from the Upper Bathonian, Middle Jurassic of Kirtlington, Oxfordshire, England. Marmorerpeton is more primitive than any other known salamander in the absence of intravertebral spinal nerve foramina in the atlantal centrum, but in other features it resembles members of the family Scapherpetontidae, neotenous salamanders otherwise known from the Upper Cretaceous and Palaeocene. The Kirtlington herpetofauna is a unique freshwater assemblage of Middle Jurassic small amphibians and reptiles, several of which represent the earliest known occurrences of their respective groups.     

Animal behavior frozen in time: gregarious behavior of Early Jurassic lobsters within an ammonoid body chamber

Direct animal behavior can be inferred from the fossil record only in exceptional circumstances. The exceptional mode of preservation of ammonoid shells in the Posidonia Shale (Lower Jurassic, lower Toarcian) of Dotternhausen in southern Germany, with only the organic periostracum preserved, provides an excellent opportunity to observe the contents of the ammonoid body chamber because this periostracum is translucent. Here, we report upon three delicate lobsters preserved within a compressed ammonoid specimen of Harpoceras falciferum. We attempt to explain this gregarious behavior. The three lobsters were studied using standard microscopy under low angle light. The lobsters belong to the extinct family of the Eryonidae; further identification was not possible. The organic material of the three small lobsters is preserved more than halfway into the ammonoid body chamber. The lobsters are closely spaced and are positioned with their tails oriented toward each other. The specimens are interpreted to represent corpses rather than molts. The lobsters probably sought shelter in preparation for molting or against predators such as fish that were present in Dotternhausen. Alternatively, the soft tissue of the ammonoid may have been a source of food that attracted the lobsters, or it may have served as a long-term residency for the lobsters (inquilinism). The lobsters represent the oldest known example of gregariousness amongst lobsters and decapods in the fossil record. Gregarious behavior in lobsters, also known for extant lobsters, thus developed earlier in earth's history than previously known. Moreover, this is one of the oldest known examples of decapod crustaceans preserved within cephalopod shells.     

Aspects of late triassic palynology. 3. Palynology of latest triassic and earliest jurassic deposits of the northern limestone alps in Austria and southern Germany,with special reference to a palynological characterization of the Rhaetian stage in Europe

The present paper provides a palynological analysis of samples from the “Kössener Schichten”, “Schattwalder Schichten” and the ammonoid (Psiloceras planorbe)-bearing beds of the “Fleckenmergel/Fleckenkalk” found in the vicinity of Salzburg (Austria and adjacent part of Germany).It is shown that the compositional development of latest Triassic—earliest Jurassic palynological assemblages from Alpine sequences matches the situation in the Germanic realm. A characteristic phase in this development is interpreted in terms of the Rhaetipollis germanicus Assemblage Zone. It is considered that this zone could well be regarded as a palynological characterization of the Rhaetian Stage in Europe and North America.Although palynology does not support the Rhaetian concept based on the occurrence of the species Choristoceras marshi exclusively, the traditional ammonoid-based Triassic—Jurassic boundary can be palynologically recognized.     

Peltoceratinae (Ammonoidea) from the upper Callovian–lower Oxfordian of Kutch,India: Systematic revision and biostratigraphic implication

The ammonoid subfamily Peltoceratinae is a well-constrained group restricted to the Middle Upper Jurassic; it had a near global distribution. In Kutch (India), they were the most important marine macroinvertebrate group during the upper Callovian–lower Oxfordian. Previous reports suffered from excessive splitting due to failure to account for the large intraspecific variability and sexual dimorphism of these ammonoids. The present systematic revision, based on numerous specimens including many near-complete adult specimens, enables to recognize and describe three species of the genus Peltoceras (P. athleta, P. ponderosum, and P. kumagunense) and two species of the genus Peltoceratoides (P. semirugosus and P. propinquus), as well as identifying their macroconchs and microconchs. We rediscovered Peltoceras athleta, which was previously described by Waagen (1875) but often overlooked by subsequent workers. We were able to identify several morphs within this species that are stunningly similar with those previously reported from France. The presence of P. athleta sensu stricto prompted us to reinstate the Athleta Zone in Kutch, improving intercontinental biostratigraphic correlation. Also, it highlights that the genus Peltoceras has a different biostratigraphic longevity in Kutch: in Europe it is restricted to the upper Callovian, whereas in India it also occurs in the lower Oxfordian.     

A new ammonoid fauna from the Carnian (Upper Triassic) Kasimlar Formation of the Taurus Mountains (Anatolia,Turkey)

We describe a new ammonoid fauna from the Taurus Mountains of southern Turkey. The Carnian ammonoid fauna from A?a?iyaylabel is presented for the first time. Ammonoids were obtained from limestone to marl beds of an approximately 35‐m‐thick section, which presents the rare opportunity to investigate ammonoid faunas across the Lower–Upper Carnian boundary. Intense sampling near the village of A?a?iyaylabel led to the recognition of a new Lower Carnian (Julian 2) to Upper Carnian (Tuvalian 1) ammonoid fauna from the Kasimlar Formation. The genus Kasimlarceltites gen. nov. is reported for the first time from the Taurus Mountains, which represents the main faunal element and occurs as huge mass occurrence (n ? 1 million). Kasimlarceltites krystyni gen. et sp. nov., Klipsteinia disciformis sp. nov. and Anasirenites crassicrenulatus sp. nov. occur within the Lower Carnian Carbonate member (Units A–B) of the Kasimlar Formation from the Taurus Platform Units. Ammonoids described from the marls of the Tuvalian Marlstone member were deposited during a major, Tethyan‐wide climate crisis – the so‐called Carnian crisis – characterized by a demise of carbonate platforms. Based on the biostratigraphic relevance of certain ammonoid taxa described herein, the age of the analysed parts of the Kasimlar Formation is Julian 2 to Tuvalian 1. The discovery of the new ammonoid assemblages from A?a?iyaylabel substantiates the significance of Upper Triassic faunas within the Taurus Mountains and facilitates the correlation with faunal assemblages from other regions in the Tethyan Realm. The ammonoid fauna and facies indicate a general deepening from open‐platform margins, over deeper shelf settings down to an open marine‐influenced basinal environment. The tentative habitat for Kasimlarceltites gen. nov. is a shallow platform environment to upper mid‐ramp.     

Ammonoid age control of the Early Triassic marine reptiles from Chaohu (South China)

Here we described a series of ammonoid specimens from the Early Triassic of Chaohu, South China and recognized the occurrence of Procolumbites for the first time in this area. The Procolumbites layer is about one meter above the first appearance of Chaohusaurus, indicating that the oldest Chaohusaurus is within the Procolumbites Zone of middle Spathian age. This new age constrain is significantly older than the previously suggested Subcolumbites Zone assignment (early late Spathian). To date, Chaohusaurus is the oldest known ichthyopterygian.     

Correlation of conodont and ammonoid successions across the Viséan–Serpukhovian boundary — A review of occurrences in the South Urals,Cantabrian Mountains,western Ireland and the Rhenish Mountains

Lochriea ziegleri is the conodont species that bears the highest potential to be the index for the base of Serpukhovian. Proposed phylogenetic lineages within the genus Lochriea, particularly the lineage L. nodosa–L. ziegleri, can be confirmed by the latest studies of the succession of the respective species in sections of north-western Ireland and the Rhenish Mountains of Germany; in these regions the first occurrence datum (FOD) of L. ziegleri is in the P2a zone or the comparable Neoglyphioceras suerlandense ammonoid zone. In the Cantabrian Mountains and the South Urals, the FODs of L. ziegleri are possibly located within ammonoid zones characterized by the genera Revilloceras and early representatives of Dombarites zones with falcatoid ornament.     

Latest Smithian (Early Triassic) ammonoid assemblages in Utah (western USA basin) and their implications for regional biostratigraphy,biogeography and placement of the Smithian/Spathian boundary

The late Smithian extinction represents a major event within the Early Triassic. This event generally corresponds to a succession of two, possibly three successively less diverse, cosmopolitan ammonoid assemblages, which when present, provide a robust biostratigraphic framework and precise correlations at different spatial scales. In the western USA basin, known occurrences of latest Smithian taxa are rare and until now, have only been documented from northeastern Nevada. Based on these restricted basinal occurrences, a regional zone representing the latest Smithian was postulated but not corroborated, as representative taxa had not yet been reported from outside Nevada. Here we document two new ammonoid assemblages from distant localities in northern Utah, overlying the late Smithian Anasibirites beds and characterized by the unambiguous co-occurrence of Xenoceltites subevolutus and Pseudosageceras augustum. The existence of a latest Smithian zone in the western USA basin is therefore validated, facilitating the identification of the Smithian/Spathian boundary and intra-basin correlation. This zone also correlates with the latest Smithian zone recognized from southern Tethyan basins. Additionally, these new data support other observed occurrences of Xenoceltites subevolutus throughout most of the late Smithian.     

Tithonian ammonoid biostratigraphy in eastern Himalayan Tibet

The Tithonian and Lower Berriasian sediments in the eastern Himalayas of Tibet contain an extensive sequence of ammonoid fauna. New collections in situ through the Lanongla, Pure, Gucuo and Jiapeila sections have facilitated a major revision of the ammonoid assemblages. Probably due to depositional facies segregation, the Belemnopsis galoi-bearing beds can be regarded as the oldest Tithonian sediments in which the basal Tithonian ammonoid Kossmatia is not present. The Lower Tithonian includes the Virgatosphinctes-Aulacosphinctoides and Uhligites-Aulacosphinctes assemblages; the Upper Tithonian includes the Haplophylloceras pingue, Blanfordiceras wallichi and Haplophylloceras strigile-Corongoceras-Himalayites assemblages. The Spiticeras assemblage is suggested to be from the Lowermost Berriasian. The new ammonoid assemblages at well-defined levels in Himalayan Tibet provide some crucial links for correlation with other regions of the SW Pacific domain where these ammonoid genera have been widely distributed.     

<Emphasis Type="Italic">Goniatites sphaericus</Emphasis> (Sowerby, 1814), the archetype of Palaeozoic ammonoids: a case of decreasing phenotypic variation through ontogeny

The Late Viséan (Early Carboniferous) ammonoid species Goniatites sphaericus (Sowerby, 1814) is revised with the use of the type material. With respect to conch morphology, suture line and particularly shell ornament, it is regarded as a senior synonym of the species Goniatites fimbriatus (Foord and Crick, 1897) and must therefore be re-established as an important index fossil for the ammonoid stratigraphy of the Rhenohercynian and Subvariscan shelves. The Goniatites fimbriatus Zone has been re-named Goniatites sphaericus Zone on this basis. The study of ontogenetic changes in the conch parameters shows that intraspecific variation occurs within a wide range. In the whorl profile, variation is very high in small juveniles, while in the coiling rate, intermediate growth stages are more strongly affected.     

AMMONOIDS AND THE TRIASSIC/JURASSIC BOUNDARY IN THE HIMALAYAS OF SOUTHERN TIBET

Abstract:  The sections of Germig in the Nyalam area, southern Tibet, provide a continuous exposure of ammonoid-bearing, uppermost Triassic and basal Jurassic strata. Eighteen species (three of them new) are described and illustrated: Choristoceras marshi Hauer , C. aff. marshi, C. cf. nobile Mojsisovics , C. nyalamense sp. nov., Eopsiloceras germigense sp. nov. , Pleuroacanthites aff. biformis (Sowerby) , Rhacophyllites sp., Nevadaphyllites cf. psilomorphus (Neumayr), Neophyllites sp. indet., Neophyllites cf. biptychus (Lange), Psiloceras tibeticum sp. nov., P. calliphyllum (Neumayr), Euphyllites cf. struckmanni (Neumayr), Discamphiceras pleuronotum (Canavari), Alsatites spp., Kammerkarites frigga , and K. sp. The ammonoid fauna shows a strong affinity to that of the Northern Calcareous Alps, although diversity in the Calliphyllum Zone is markedly lower. The ammonoid succession across the Triassic/Jurassic boundary is subdivided into four zones: the Rhaetian Marshi, the basal Hettangian Tibeticum, the lower Hettangian Calliphyllum, and the middle Hettangian Pleuronotum zones. It is the only known succession across the Triassic/Jurassic boundary in the Tethyan Realm that is not condensed. The Marshi and Calliphyllum zones are correlated with the same zones in the Northern Calcareous Alps. The Tibeticum Zone, a new local zone, is transitional between the Marshi and the Calliphyllum zones in that it yields both choristoceratids and psiloceratids. Its base is taken to mark the base of the Jurassic System in the eastern Tethys.     

Desmoceras (Pseudouhligella) intrapunctatum n. sp. (Ammonoidea) aus dem Unter-Albium von NW-Madagaskar mit Ritzstreifen

The new ammonoid speciesDesmoceras (Pseudouhligella) intrapunctatum (DesmoceratoideaZittel, 1895) is described and figured from the dark glauconitic marls of the Lower Albian of Ambatolafia (Mahajanga Basin of northwestern Madagascar). On the molds of its body chamber a well developed system of dotted lines is visible originally described from Palaeozoic ammonoids and nautiloids (= ?Ritzstreifen“Sandberger & Sandberger 1850 sensuTozer 1972). These internal shell structures have been originated by small ridges of the inner prismatic layer as the result of an incomplete mineralization. “Ritzstreifen” could not have observed in any other ammonoid species accompanied withDesmoceras (Pseudouhligella) intrapunctatum, despite of the same excellent aragonitic shell preservation. Therefore, this unique character is interpreted as taxonomically significant on species level.     

Ecpagloxylon mathiesenii gen. nov. et sp. nov., a Jurassic wood from Greenland with several primitive angiosperm features

Fossil wood specimens from the late Early–early Middle Jurassic of Jameson Land, Eastern Greenland, have several unexpected features: tracheids of irregular size and shape, thinly pitted ray cell walls, heterogeneous rays, partially scalariform radial pitting, both areolate and simple pits, and pitted elements associated with rays. These characters diverge markedly from those typical of Jurassic wood, which usually conform to those of modern conifers. Although this combination of features is not encountered in any extant angiosperm, each has been documented in one or several extant homoxylous angiosperms, particularly Amborella, Trochodendron, and Tetracentron. As these wood specimens are not found in connection with any reproductive part, it is impossible to confidently assign them to the angiosperms. If a Jurassic angiosperm did exist, however, it might well have had a similar wood. This material is an early bench-mark in the evolution that led from homoxylous conifer-like wood to that of the angiosperms. Its particular biogeography (Arctic) could renew the discussion about the area of origin of the angiosperms.     

Ammonoid recovery from the Late Permian mass extinction event

Previous research indicated that ammonoid taxonomic diversity exploded after the Late Permian mass extinction, regaining pre-extinction levels by the Late Induan (Dienerian substage). From taxonomic analyses it had been inferred that ammonoids recovered rapidly, relative to other marine invertebrate groups. Complementing taxonomic metrics with morphologic and spatial data revealed more complex recovery dynamics. Morphological analysis indicated that ammonoids did not fully recover until the Spathian or Anisian. Taxonomic diversity is a poor predictor of disparity during the recovery. Spatial partitioning of taxonomic and morphological diversity revealed spatially homogeneous recovery patterns. Combining taxonomic, morphological, and spatial data refined interpretations of Triassic ammonoid recovery patterns and indicated that ecological, not intrinsic, factors were the probable control on ammonoid recovery rates. To cite this article: A.J. McGowan, C. R. Palevol 4 (2005).     

Shell Microstructure of <Emphasis Type="Italic">Rhynchonella loxiae</Emphasis> Fischer (Brachiopoda,Rhynchonellida) from the Upper Volgian of Moscow Region

The shell microstructure of Late Jurassic rhynchonellids of the family Rhynchonellidae from the boreal basin of the Russian Platform is studied for the first time. The studied rhynchonellids differ in the shell microstructure from the Late Jurassic and Early Cretaceous rhynchonellids of the families Cyclothyrididae, Praecyclothyrididae, Basiliolidae, and Norellidae from the warm-water Mediterranean basin. The revealed peculiarities of shell structure of Rhynchonella loxiae (type species of Rhynchonella) may be added to the diagnosis of Rhynchonella and should be considered in characterizing the family Rhynchonellidae.     

Iterative ontogenetic development of ammonoid conch shapes from the Devonian through to the Jurassic

We measured longitudinal growth in conch cross‐sections of 177 Devonian to Jurassic ammonoid species to test whether conch ontogenetic development parallels the iterative evolution of pachyconic or globular conch shapes. Ontogenetic trajectories of two cardinal conch parameters, conch width index and umbilical width index, show a few common recurring ontogenetic pathways in terms of the number of ontogenetic phases. The most common, with three phases in the conch width index (decrease–increase–decrease) and umbilical width index (increase–decrease–increase), is termed here C‐mode ontogeny (after the Carboniferous genus Cravenoceras). Many of the studied globular Palaeozoic and Triassic species (of the latter, particularly the arcestid ammonoids) share principal patterns in the triphasic C‐mode conch ontogeny in closely related groups but also between unrelated groups as well. The repetition of conch growth patterns is an example of convergent evolution of the entire life history of globular ammonoids. The studied Jurassic globular shaped ammonoids deviate from the growth patterns seen in earlier groups showing less pronounced ontogenetic trajectories with nearly isometric or weakly asymmetric growth without distinct phases. This trajectory is termed here M‐mode ontogeny (after the Jurassic genus Macrocephalites). No major change in the ontogenetic modes of pachyconic and globular ammonoids occurred moving from the Palaeozoic into the Mesozoic; the survivors of the end‐Permian extinction event iteratively developed conch ontogenies similar to those of Palaeozoic forms. In contrast, the Triassic–Jurassic boundary marks the major event with the evolution of some cardinal conch parameters relating to globular ammonoid ontogeny.