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.     

Large‐scale evolutionary trends of Acrochordiceratidae Arthaber, 1911 (Ammonoidea,Middle Triassic) and Cope’s rule

Abstract: Directed evolution of life through millions of years, such as increasing adult body size, is one of the most intriguing patterns displayed by fossil lineages. Processes and causes of such evolutionary trends are still poorly understood. Ammonoids (externally shelled marine cephalopods) are well known to have experienced repetitive morphological evolutionary trends of their adult size, shell geometry and ornamentation. This study analyses the evolutionary trends of the family Acrochordiceratidae Arthaber, 1911 from the Early to Middle Triassic (251–228 Ma). Exceptionally large and bed‐rock‐controlled collections of this ammonoid family were obtained from strata of Anisian age (Middle Triassic) in north‐west Nevada and north‐east British Columbia. They enable quantitative and statistical analyses of its morphological evolutionary trends. This study demonstrates that the monophyletic clade Acrochordiceratidae underwent the classical evolute to involute evolutionary trend (i.e. increasing coiling of the shell), an increase in its shell adult size (conch diameter) and an increase in the indentation of its shell suture shape. These evolutionary trends are statistically robust and seem more or less gradual. Furthermore, they are nonrandom with the sustained shift in the mean, the minimum and the maximum of studied shell characters. These results can be classically interpreted as being constrained by the persistence and common selection pressure on this mostly anagenetic lineage characterized by relatively moderate evolutionary rates. Increasing involution of ammonites is traditionally interpreted by increasing adaptation mostly in terms of improved hydrodynamics. However, this trend in ammonoid geometry can also be explained as a case of Cope’s rule (increasing adult body size) instead of functional explanation of coiling, because both shell diameter and shell involution are two possible paths for ammonoids to accommodate size increase.     

New aspects in ammonoid mode of life and their distribution

The intensively debated functional morphology and mode of distribution of ammonites can be clarified and explained when ammonoids are regarded as conch-bearing octopods. The terminal body chambers of some ammonites were modified into a floating egg case, widely dispersing the hatchlings along the course of oceanic and long-shore currents. Hatchlings from eggs attached to a substrate lived and bred in the same region, developing indigenous evolutionary lineages. Females became sexually mature after 1–3 years of age, breeding only once, dispatching numerous eggs at a time. This contributed to the high evolutionary rate of ammonoids. Due to ammonoid short longevity, growth was rapid and septa were frequently precipitated. Ammonite internal molds exhibit small scars of adductor muscles, which could rapidly detach and reattach during septa secretion. The resultant weak hold between the conch and the body was compensated by the septal marginal fluting in the form of backward expanding lobes, into which the soft tissue penetrated, stiffening when needed. Increased suture complexity (unrelated to buoyancy regulation or diving ability) reflects a better hold between the body and the buoyant conch, hence a more successful functioning. The complex network of mantle muscle fibers could also form the template for septa precipitation. The high intelligence and learning ability of extant octopods can explain ammonoids’ adaptation to diverse niches, successfully coping with ecological changes and threats (hence evolution) in contrast to the associated nautiloids. Post-mortal drift of the empty conch was minor due to rapid sinking of shells of dead ammonoids, for which ammonites are good biogeographic indicators.     

AMMONOIDS ACROSS THE PERMIAN/TRIASSIC BOUNDARY: A CLADISTIC PERSPECTIVE

Abstract:  The rapid diversification of ceratitid ammonoids during the earliest Mesozoic has been taken at face value as an example of explosive radiation following the Permian/Triassic mass extinction. However, the validity of this interpretation has never been tested within a phylogenetic framework. A total evidence cladistic analysis of Mid–Late Permian and Induan (earliest Triassic) ammonoids confirms the monophyly of the Ceratitida. Partitioned phylogenetic analysis of suture line characters vs. shell shape and ornament characters confirms the importance of suture-line characters for resolving the higher taxonomy of ammonoids. When the cladogram is compared with the observed fossil record, the resultant tree implies that the divergence of a number of early Triassic lineages actually occurred during the latest Permian. If these range extensions are taken into account the ammonoid per-genus extinction rate across the Permian/Triassic boundary drops from c. 85 per cent to c. 60 per cent.     

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.     

High-resolution biochronology and diversity dynamics of the Early Triassic ammonoid recovery: The Smithian faunas of the Northern Indian Margin

Based on new collections of abundant and well preserved material from the Salt Range (Pakistan), Spiti (Northern India) and Tulong (South Tibet), several recent studies focused on the taxonomic revision and detailed biostratigraphy of Smithian ammonoids. In this work, biochronological data for these three well-documented basins are analyzed by means of the Unitary Associations method, resulting in a biochronological scheme of unprecedented high-resolution for the Smithian of the Northern Indian Margin (NIM). Data for each basin are first processed separately, thus yielding three local biochronological zonations. Then, the three sequences are processed together as a regional three-section data set for the construction of an inter-basin sequence at the NIM level. The latter zonation comprises 16 Unitary Associations grouped into 13 zones for the entire Smithian. Analysis of ammonoid diversity dynamics based on this new highly resolved time frame highlights (i) a marked diversification during the early Smithian, (ii) a severe extinction during the late Smithian, and (iii) an overall very high turnover throughout the Smithian. At a global spatial scale and stage resolution, the diversity of Smithian ammonoid genera appears surprisingly high, as highlighted by a previous study. It is shown that at a smaller geographic scale and with the most highly resolved time frame, Smithian ammonoids of the NIM reached their explosive diversity peak essentially through extremely high turnover rates rather than through a classic diversification process of high origination rates coupled with low extinction rates. Based on recently published U/Pb ages, regional apparent total rates of origination and extinction of more than 100 species per My can be inferred for the Smithian ammonoids of the NIM.     

Allometric growth and intraspecific variability in the basal Carboniferous ammonoid<Emphasis Type="Italic">Gattendorfia crassa</Emphasis><Emphasis Type="SmallCaps">Schmidt</Emphasis>, 1924

Gattendorfia crassa is an Early Carboniferous (Mississippian) goniatite species with strikingly allometric conch growth. Analysis of 15 high-precision cross-sections of this species demonstrates the small intraspecific variability of some of the conch form characters, but remarkable variability in others. While the whorl expansion rate, umbilical width, and conch thickness vary within narrow limits, the expansion rates of the whorl height and whorl width are remarkably plastic. Variability of most of the characters tends to be smallest in intermediate growth stages, whereas juveniles and adults are more variable. The differences in morphological plasticity are interpreted in terms of the function of the ammonoid conch, especially the orientation of the aperture during life.      

Major trends in the evolution of Permian ammonoids

The phylogeny of major families of Permian ammonoids is analyzed. The evolution of most families followed a typical scenario with distinct stages of early evolution, diversification, and decline. A smaller group followed a different evolutionary narrative, with indistinct stages. The former group includes families with both simple and complex morphology and a wide range of variation. The nature and trends in the evolution of the families may change depending on their phylogenetic stage. The Early Permian (Asselian), the second half of the Artinskian, and the beginning of the Middle Permian were marked by the most significant evolutionary changes. The Late Permian was the time of the decline of Paleozoic ammonoid orders and of the onset of the evolution of the Mesozoic order Ceratitida.     

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).     

Early evolutionary trends in ammonoid embryonic development

During the Devonian Nekton Revolution, ammonoids show a progressive coiling of their shell just like many other pelagic mollusk groups. These now extinct, externally shelled cephalopods derived from bactritoid cephalopods with a straight shell in the Early Devonian. During the Devonian, evolutionary trends toward tighter coiling and a size reduction occurred in ammonoid embryonic shells. In at least three lineages, descendants with a closed umbilicus evolved convergently from forms with an opening in the first whorl (umbilical window). Other lineages having representatives with open umbilici became extinct around important Devonian events whereas only those with more tightly coiled embryonic shells survived. This change was accompanied by an evolutionary trend in shape of the initial chamber, but no clear trend in its size. The fact that several ammonoid lineages independently reduced and closed the umbilical window more or less synchronously indicates that common driving factors were involved. A trend in size decrease of the embryos as well as the concurrent increase in adult size in some lineages likely reflects a fundamental change in reproductive strategies toward a higher fecundity early in the evolutionary history of ammonoids. This might have played an important role in their subsequent success as well as in their demise.     

滇东南泸西县上三叠统卡尼阶小凹组菊石*

本文对云南省东南部的泸西县白水镇小兴安村上三叠统卡尼阶小凹组菊石类进行了详细研究,共发现菊石9属13种,其中Anasirenites属为华南首次发现。依据各物种组合特征及其地层分布情况,自下而上识别出菊石Trachyceras multituberculatum带和Austrotrachyceras triadicum带,时代对应于晚三叠世早卡尼期的Julian 1和Julian 2。该菊石组合不仅记录了卡尼期Julian期Trachyceratidae的辐射事件,同时该菊石组合与卡尼期特提斯域和泛大洋域的菊石组合均有良好的对比性,并且为华南古地理研究提供了新的补充材料。另外,通过对剖面的岩性特征与菊石生物地层特征分析,认为在滇东南地区由于卡尼期湿润幕事件(CPE)所导致的碳酸盐岩台地消亡起始时间应为Julian 1末期略早于陆源碎屑输入的Julian 2亚期。     

Morphospace occupation of ammonoids over the Devonian-Carboniferous boundary

In a case study, a stratophenetic analysis of the succeeding ammonoid faunas of the latest Devonian and earliest Carboniferous cephalopod limestones of the Rhenish Massif has been made. This investigation concentrated on the development of the whorl expansion rate (WER), a character very important for ammonoids since it indicates the body chamber length and hence the orientation in the water column as well as mobility. The study leads to the conclusion that the Hangenberg Event caused an almost complete change in the morphospace adopted by ammonoids. All clymenüds as well as tornoceratids became extinct at or immediately after the Hangenberg Event, and the morphospace left behind by these Devonian groups was reoccupied only incompletely by the surviving prionoceratid ammonoids.     

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.     

First direct evidence of ammonoid ovoviviparity

Ammonoids had high evolutionary rates and diversity throughout their entire history and played an important role in the high‐resolution sub‐division of the Mesozoic, but much of their palaeobiology remains unclear, including the brooding habitat. We present our study of the first recorded ammonite embryonic shell clusters preserved with calcified embryonic aptychi in situ within the body chambers of mature macroconch shells of the Early Aptian (Early Cretaceous) ammonite Sinzovia sazonovae. The following support the idea that the clusters are egg masses, which developed inside ammonite body chambers: the absence of post‐embryonic shells and any other fossils in these clusters, the presence of the aptychi in all embryonic shell apertures and peculiarities of adult shells preservation. These facts confirm earlier speculations that at least some ammonoids could have been ovoviviparous and that, like many modern cephalopods, they could have reproduced in mass spawning events. The aptychi of ammonite embryonic shells are observed here for the first time, indicating that they were already formed and calcified before hatching. Our results are fully congruent with the peculiar modes of ammonoid evolution: quick recovery after extinctions, distinct evolutionary rates, pronounced sexual dimorphism and the nearly constant size of embryonic shells through ammonoid history. We assume that adaptation to ovoviviparity may be the reason for the presence of these features in all post‐Middle Devonian ammonoids.     

Nautilus—a poor model for the function and behavior of ammonoids?

Inferences drawn from the biology, function, and behavior of closely related living forms facilitate interpretation of the mode of life of groups known only from the fossil record. The choice of phylogenetically relevant modern 'model organisms' can have critical bearing on the resulting interpretations. The biology and behavior of fossil ammonoids are often interpreted in the light of evidence derived from the study of modern Nautilus . However, examination of the fossil record and cladistic analyses both indicate that coleoids are much more closely related to ammonoids than is Nautilus . Coleoid biology and behavior differ dramatically from the biology and behavior of Nautilus . Thus, the inclusion of coleoids as examples, rather than reliance on Nautilus alone, produces a strikingly different vision of ammonoid biology and suggests that inferences of ammonoid biology and behavior that rely exclusively on Nautilus should be reviewed. Two features related to swimming ability in Nautilus , static stability and large retractor muscles, are much reduced in many ammonoids, leading to the interpretation that ammonoids were poorer swimmers than Nautilus . However, reexamination of the evidence indicates that static stability should not play a role in the swimming of ammonoids with long body chambers. In addition, functional arguments suggest that a coleoid-like swimming mechanism should have evolved prior to the loss of the body chamber in coleoids. Thus, a coleoid-like swimming mechanism is likely to have evolved prior to the separation of ammonoid and coleoid lineages. A mechanism is proposed by which a coleoid swimming mechanism, independent of retractor muscle size, could function in ammonoids with long body chambers.□ Ammonoids, ammonites, evolution, functional morphology , Nautilus, phylogeny .     

Recent advances in heteromorph ammonoid palaeobiology

Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non-planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well-developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus-type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U-shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near-neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near-horizontal aperture pointing upwards. Heteromorphs with a U-shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep-subtidal to offshore facies but are rare in shallow-subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep-subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow-subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro- and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40–500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.     

On the origin of bactritoids (Cephalopoda)

There is a high probability that bactritoids represent a paraphylum or polyphylum. The initial chambers or protoconchs of the Early-Middle DevonianBactrites Sandberger,Devonobactrites Shimansky, andLobobactrites Schindewolf are elongated spheres with a diameter of 0.3–1.0 mm. The initial chambers are larger in diameter than the slender, smooth shaft located adorally to the initial chamber. Similar apices occur in a number of Late Silurian sphaerorthoceridans with central siphuncles. Sphaerorthoceridans with a bactritoid-like apex and an eccentric siphuncle are known from the Early Devonian. The earliest questionableBactrites occurs in the Pragian (middle Early Devonian). By Emsian time bactritoids are common elements of cephalopod faunas.Bactrites-like orthocones of the Middle Ordovician and Late Silurian are homeomorphs with clearly different early growth stages. Thus, the time interval between the first appearance ofBactrites and the origin of ammonoids can be narrowed down to the Pragian to Early Emsian. The placement of the siphuncle in a ventral marginal position has been used as one of the critical morphologic features in defining the bactritoids. However, the displacement of the siphuncle from subcentral or eccentric positions toward the conch margin occurred at least three times during the Ordovician — Early Devonian evolution of the Orthocerida. Thus, there is a high probability that a marginal shift of the orthocerid siphuncle occurred in post-Emsian times, too.     

Ontogeny and evolution within the Myophorellidae (Bivalvia): Paedomorphic trends

The role of heterochronic phenomena in molluscan evolution is insufficiently understood but potentially significant. The aim of this paper is to explore some paedomorphic trends in the evolution of the Myophorellidae (Bivalvia: Trigoniida). Early ontogeny of general shell shape and ornamentation of one species of Steinmanella was analyzed and compared to data obtained for three species of Myophorella: two belonging to the subgenus M. (Promyophorella) (one from the Jurassic and one from the Cretaceous) and one belonging to the Jurassic M. (Myophorella). For general shell shape, a geometric morphometric analysis was performed on lateral views of the shells. Regarding ornamentation, flank costal disposition on the marginal carina, tubercle separation and relative development of the sub-commarginal subset of flank costae were quantified. A qualitative analysis was also performed. A two-trend shell shape development is considered as primitive. The first trend is marked by a relative reduction of the posterior margin together with a relative elongation of the shell. A tangential opisthogyrate growth component characterizes the second trend. There is a transitional stage where both trends interact. Early flank ornamentation is characterized by two or three sub-commarginal costae, continuous through the area, after which oblique costae with fine tubercles start to form. The subgenus M. (Myophorella) evolved by paedomorphic retention of juvenile shell shape and ornamentation, resulting in a large shell with coarse tubercles. Shell morphology in Steinmanella evolved by paedomorphic suppression of the primitive second trend in the development of the shell, resulting in an orthogyrate shell shape, and the retention of juvenile ornamentation (coarse tubercles, more sub-commarginal costae, juvenile rates of costal disposition). The paedomorphic (most likely by deceleration) retention of juvenile shell morphology within the Myophorellidae seems to have been recurrent within the group, resulting in many cases of convergence, and obscuring the phylogenetic relationships among its species.     

Occluded Umbilicus In The Pinacitinae (Devonian) And Its Palaeoecological Implications

Eifelian (Middle Devonian) ammonoids of the Pinacitinae Hyatt, 1900 ( Exopinacites , Pinacites )with preserved shell structures from the eastern Anti–Atlas (Morocco) have revealed unusual morphological features. The Pinacitinae belong to the earliest ammonoids which closed their umbilici. As an approach to an interpretation of these structures, the representatives of the subfamily Pinacitinae ( Exopinacites singularis , Pinacites jugleri , P. eminens ) are compared with other ammonoids, e.g. Acrimeroceras , Araucanites , Clistoceras , Gaudryceras , Nathorstites , Prolobites , and Synpharciceras , which produced umbilical plugs and covers. Some of these are comparable in structure to Nautilus pompilius and N. belauensis . In contrast to all of these taxa, the lateral shell wall of the Pinacitinae reached the centre of the umbilicus and formed an umbilical lid. The umbilical shell wall rests on the umbilical lid of the previous whorl. This construction probably had the advantage that it improved the hydrodynamic properties of the conch, along with the oxyconic conch shape and the approximately horizontal orientation of the aperture.     

The two Early Toarcian (Early Jurassic) extinction events in ammonoids

The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass‐extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass‐extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass‐extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long‐ranging ammonoids) and Polyplectus (relatively long‐ranging compared to other Ammonitina).