Untangling phylogenetic,geometric and ornamental imprints on Early Triassic ammonoid biogeography: a similarity‐distance decay study

Ammonoids are diverse and widespread fossil, externally shelled cephalopods that flourished for more than 300 Myr before their total extinction 65 Ma ago. In spite of two centuries of intensive scientific studies, their mode(s) of life and long‐distance dispersal abilities remain poorly known. Here, we address this by focusing on the latitudinal distribution of Early Triassic (approximately 250 Myr) ammonoids through similarity‐distance decay analyses. We examine and compare rates of similarity‐distance decay between various groups with respect to systematics, shell geometry and ornamentation to untangle phylogenetic, geometric and ornamental imprints on the observed biogeographical pattern. Our data do not support any phylogenetic and shell ornamentation influence, but rather demonstrate the significant effect of (sub‐)adult shell geometry on the similarity–distance decay: most evolute morphs tend to have been more endemic than most involute forms. This contrasts with the classic hypothesis that long‐distance ammonoid dispersal mainly occurred during the earliest planktonic stages, and thus that (sub‐)adult morphological characteristics should not constrain large‐scale biogeographical patterns of ammonoids. Although direct control by Sea Surface Temperature can be discarded, this result may indicate that at least some adult Triassic ammonoid morphs were skilled active swimmers capable of achieving long‐distance migration, as observed for some present‐day coleoid cephalopods. □Ammonoid, dispersal, similarity‐distance decay, morphology, phylogeny, biogeography, Triassic.     

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.     

Revision of the genus Acrochordiceras Hyatt, 1877 (Ammonoidea,Middle Triassic): morphology,biometry, biostratigraphy and intra‐specific variability

Abstract: The family Acrochordiceratidae Arthaber, 1911 ranges in age from latest Spathian to the middle/late Anisian boundary, and it represents a major component of ammonoid faunas during that time. The middle Anisian genus Acrochordiceras Hyatt, 1877 is the most widespread taxon of the family and occurs abundantly worldwide within the low paleolatitude belt. However, there is a profusion of species names available for Acrochordiceras. This excessive diversity at the species level essentially results from the fact that sufficiently large samples were not available, thus leading to a typological approach to its taxonomy. Based on new extensive collections obtained from the Anisian (Middle Triassic) Fossil Hill Member (Star Peak Group, north‐west Nevada) for which a high resolution biostratigraphic frame is available, the taxonomy and biostratigraphy of the genus Acrochordiceras Hyatt, 1877 is herein revised with respect to its intra‐specific variation. Morphological and biometric studies (c. 550 bedrock‐controlled specimens were measured) show that only one species occurs in each stratigraphic level. Continuous ranges of intra‐specific variation of studied specimens enable us to synonymize Haydenites Diener, 1907, Silesiacrochordiceras Diener, 1916 and Epacrochordiceras Spath, 1934 with Acrochordiceras Hyatt, 1877. Three stratigraphically successive species are herein recognized in the low paleolatitude middle Anisian faunas from Nevada: A. hatschekii (Diener, 1907), A. hyatti Meek, 1877 and A. carolinae Mojsisovics, 1882. Moreover, an assessment of intra‐specific variation of the adult size range does not support recognition of a dimorphic pair (Acrochordiceras and Epacrochordiceras) as previously suggested by other workers (Epacrochordiceras is the compressed and weakly ornamented end‐member variant of Acrochordiceras). The successive middle Anisian species of Acrochordiceras form an anagenetic lineage characterized by increasing involution, adult size and intra‐specific variation. This taxonomic revision based on new bedrock‐controlled collections is thus an important prerequisite before studying the evolution of the group.     

From near extinction to recovery: Late Triassic to Middle Jurassic ammonoid shell geometry

The Triassic–Jurassic extinction resulted in the near demise of the ammonoids. Based on a survey of ammonoid expansion rates, coiling geometry and whorl shape, we use the Raup accretionary growth model to outline a universal morphospace for planispiral shell geometry. We explore the occupation of that planispiral morphospace in terms of both breadth and density of occupation in addition to separately reviewing the occurrence of heteromorphs. Four intervals are recognized: pre‐extinction (Carnian to Rhaetian); aftermath (Hettangian); post‐extinction (Sinemurian to Aalenian) and recovery (Bajocian to Callovian). The pre‐extinction and recovery intervals show maximum disparity. The aftermath is marked by the disappearance of heteromorphs and a dramatic reduction in the range of planispiral morphologies to a core area of the morphospace. It is also characterized by an expansion into an evolute, slowly expanding part of the morphospace that was not occupied prior to the extinction and is soon abandoned during the post‐extinction interval. Aftermath and post‐extinction ammonoid data show a persistent negative correlation whereby rapid expansion rates are associated with narrow umbilical widths and often compressed whorls. The permanently occupied core area of planispiral morphospace represents generalist demersals whose shells were probably optimizing both hydrodynamic efficiency and shell stability. All other parts of the planispiral morphospace, and the pelagic modes of life the shells probably exploited, were gradually reoccupied during the post‐extinction interval. Planispiral adaptation was by diffusion away from the morphospace core rather than by radical jumps. Recovery of disparity was not achieved until some 30 Myr after the extinction event.     

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.     

Extensive intraspecific variation in a Triassic ammonoid from Siberia

The ammonoid species Czekanowskites rieberin.sp . from the Lower Anisian of Arctic Siberia (River Olenek) shows an enormous range of intraspecific variability. The 700 investigated specimens are packed in one single concretion. They are all adult with preserved living chambers and apertural structures. Morphologically they range from keeled smooth suboxycones with narrow umbilicus, through feebly ribbed platycones with a little wider umbilicus, to subcadicones with relatively wide umbilicus and straight ribs with bullae. The morphological variation is continuous; all specimmh belong to a single genetically linked population, variants of one biospecies. This phenomenon, which is quite common in boreal Triassic ammonoid faunas, has in the past led to serious taxonomic oversplitting. It presents a serious challenge to current ideas about a close correlation between modeof life and shell morphology in arnmonoids.□ Amntonoidea, boreal Triassic, ecology. Siberia, variability .     

Ammonoid palaeobiology

This, the second in the series of virtual issues of Palaeontology, charts the development of concepts in ammonoid palaeobiology, with reference to nine selected papers published in this journal from 1965 onwards. These cover a broad range of topics: goniatite bed palaeoecology, systematics and evolution in Scaphites, flow dynamics of cephalopod shells, shell growth and differential geometry, septal function, ammonoid assemblage analysis, evolutionary trends, stratocladistics, and soft part preservation. Despite the growth and success of palaeobiological interpretations of ammonoid evolution, life, morphology and environments over the last 50 years, ammonoid research retains a great potential for the future.     

Support from early juvenile Jurassic,Cretaceous and Holocene thecideoid species for a postulated common early ontogenetic development pattern in thecideoid brachiopods

Abstract: The analysis of morphological characters exhibited by the earliest ontogenetic stages of Middle Jurassic, Cretaceous and Holocene thecideoid species, together with the application of techniques that reveal the presence of early juvenile morphological characters buried in the shell fabric of adult representatives of the species concerned, reinforce the idea of a development pattern common to thecideoids from the early Middle Jurassic to the present day. Basic and long‐standing differences in the architecture of the brachidial skeleton of thecidellinids and lacazellins may be correlated with the morphology of thecospiroids and indicate that the Thecidellinidae and Thecideidae may have emerged as sister groups as early as the Triassic.     

Growth trajectories of some major ammonoid sub‐clades revealed by serial grinding tomography data

Molluscs such as ammonoids record their growth in their accretionary shells, making them ideal for the study of evolutionary changes in ontogeny through time. Standard methods usually focus on two‐dimensional data and do not quantify empirical changes in shell and chamber volumes through ontogeny, which can possibly be important to disentangle phylogeny, interspecific variation and palaeobiology of these extinct cephalopods. Tomographic and computational methods offer the opportunity to empirically study volumetric changes in shell and chamber volumes through ontogeny of major ammonoid sub‐clades in three dimensions (3‐D). Here, we document (1) the growth of chamber and septal volumes through ontogeny and (2) differences in ontogenetic changes between species from each of three major sub‐clades of Palaeozoic ammonoids throughout their early phylogeny. The data used are three‐dimensional reconstructions of specimens that have been subjected to grinding tomography. The following species were studied: the agoniatitid Fidelites clariondi and anarcestid Diallagites lenticulifer (Middle Devonian) and the Early Carboniferous goniatitid Goniatites multiliratus. Chamber and septum volumes were plotted against the septum number and the shell diameter (proxies for growth) in the three species; although differences are small, the trajectories are more similar among the most derived Diallagites and Goniatites compared with the more widely umbilicate Fidelites. Our comparisons show a good correlation between the 3‐D and the 2‐D measurements. In all three species, both volumes follow exponential trends with deviations in very early ontogeny (resolution artefacts) and near maturity (mature modifications in shell growth). Additionally, we analyse the intraspecific differences in the volume data between two specimens of Normannites (Middle Jurassic).     

The first scallop

Scallops (Pectinidae) are a highly diverse bivalve family with a long evolutionary history, but insufficient knowledge on the internal shell characters of initial taxa has hampered clarification of their phylogenetic roots. Here, morphological details of the shell interior of the basal pectinid Pleuronectites laevigatus from the Middle Triassic are documented for the first time. It is shown that ligament morphology, lack of internal buttresses and hinge articulation, presence of a right anterior auricular scroll, procrescent discs, and differential valve convexity of Pleuronectites link Pectinidae with Early Triassic aviculopectinoids rather than with entoliids, as recently proposed. The key adaptations of Pectinidae, i.e. the ctenolium and the alivincular-alate ligament system, probably evolved in conjunction with a marked size increase that required improvements in the shell attachment and in the system for opening the valves. Although Pleuronectites laevigatus is recognized as the earliest known member of the Pectinidae, a diphyletic origin of this family from different stocks within the Aviculopectinoidea cannot be ruled out.     

Diversity and palaeoecology of a highly diverse Late Triassic marine biota from the Cassian Formation of north Italy

A sample of marine invertebrates from the Late Triassic Cassian Formation (north Italy) yielded one of the most diverse Early Mesozoic fossil assemblages ever reported (c. 170 species). The assemblage was found in basin clays, but was transported from nearby carbonate platforms as indicated by fragmentation, microbial encrustation and the presence of coated grains and ooids. Most of the specimens are small (< 1 cm) reflecting both, small adult sizes and size sorting during transport. Rarefaction analysis suggests that diversity of surface collection and bulk sampling is the same. However, rank abundance, species richness and taxonomic composition differ strongly according to sampling method. Low‐grade lithification of the sediments is the main reason that high diversity can be recognized, because it facilitates disaggregation and finding of small molluscs. Sample standardization shows that the studied assemblage is much more diverse than known Early Triassic assemblages. However, its diversity is similar to that of Anisian assemblages. This suggests that recovery from the end‐Permian mass‐extinction was quite advanced in the Middle Triassic and alpha‐diversity remained high until the Late Triassic. According to current models, Early Triassic and Anisian faunas match the niche overlap phase of recovery during which diversity is built up by increasing alpha‐diversity, whereas beta‐diversity rises slowly. Subsequently, habitat width of species contracts because of increasing competition, making beta‐diversity the principal drive of overall diversity increase. The diversity pattern of various Late Triassic Cassian associations meets the predictions for the transition from the niche overlap to the habitat contraction phase.: Triassic, Cassian Formation, palaeoecology, diversity, mollusc dominance.     

Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

Appearance of morphological novelty in a hybrid zone between two species of land snail

On the small oceanic island of Chichijima, two endemic species of land snails, Mandarina mandarina and M. chichijimana, have discrete distributions separated by a hybrid zone. This study investigates the potential of hybridization as a source of morphological novelty in these snails. Mandarina mandarina possesses a shell with a higher whorl expansion rate and a smaller protoconch than M. chichijimana, relative to shell size. The number of whorls and shell size of M. mandarina do not differ from those of M. chichijimana, because the effect of higher expansion rate on number of whorls and size of the former is compensated for by its smaller protoconch. The whorl expansion rate and protoconch diameter of the individuals from the hybrid populations are intermediate or typical of either of the two species, and their average values show clinal changes along the hybrid zone. However, the hybrid populations include exceptionally high shells with many whorls and flat shells with few whorls, which are never found in the pure populations of either species. In addition, gradual increases in variance in shell height and number of whorls were found from the edges to the center of the hybrid zone. A combination of low expansion rate (typical of M. chichijimana) and a small protoconch (typical of M. mandarina) produces a shell with an extremely large number of whorls because of the geometry of shell coiling. However, the combination of high expansion rate and a large protoconch produces a shell with an extremely small number of whorls. Because of the correlation between the number of whorls and shell height, shells with an exceptional number of whorls possess an extraordinarily high or flat spire. Hybrids can inherit a mosaic of characters that, as they play out during growth, lead to novel adult morphologies. These findings emphasize the importance of hybridization as a source of morphological variation and evolutionary novelty in land snails.     

Ammonoid evolution in marine ecosystems prior to the Permian-Triassic crisis

Evolutionary trends in ammonoid communities in the Middle and Late Permian are analyzed in connection with the changes in their taxonomic, morphological, geographical, and ecological structure. It is shown that the processes that led to the almost complete disappearance of the subclass began long before the global Permian-Triassic crisis.     

Large tetrapod burrows from the Middle Triassic of Argentina: a behavioural adaptation to seasonal semi‐arid climate?

Krapovickas, V., Mancuso, A.C., Marsicano, C.A., Domnanovich, N.S. & Schultz, C.L. 2013: Large tetrapod burrows from the Middle Triassic of Argentina: a behavioural adaptation to seasonal semi‐arid climate? Lethaia, Vol. 46, pp. 154–169. We report the discovery of large burrow casts in the early Middle Triassic Tarjados Formation, at Talampaya National Park, north‐western Argentina. Facies analysis indicates the burrows are preserved in sandbars deposited by an ephemeral river under semi‐arid and seasonal climatic conditions. The structures are mostly preserved in longitudinal cross‐section and consist of an opening, an inclined tunnel (ramp), and a terminal chamber. The ramp is 8–14 cm in height, up to 130 cm in length and penetrates 49–63 cm bellow the palaeosurface with an inclination of 22°–30°. We studied burrow cast dimensions, overall architectural morphology, surficial marks, and compared them with other large burrows of both invertebrate and vertebrate origin. A tetrapod origin of the burrow casts was established based on: distinctive architecture, and size, which is more than twice the most common size range for large terrestrial invertebrate burrows. Comparison with other Upper Permian and Triassic tetrapod burrows allows us to identify three general morphological groups: (1) simple inclined burrows; (2) helical burrows; and (3) burrow network complexes, representing different behaviours. A study of tetrapod body fossils preserved within other Upper Permian and Triassic burrows shows that the Tarjados structures were most likely produced by non‐mammalian cynodonts. The environmental and climatic context suggests that aridity and seasonality played a fundamental role selecting burrowing behaviour in therapsids and that by the Early–Middle Triassic their burrowing behaviour attained a complexity comparable to modern mammals. □Argentina, behaviour, palaeoclimate, Permo‐Triassic, Tarjados Formation, Tetrapod burrows.     

Disentangling the effects of intraspecies variability,phylogeny, space,and climate on the evolution of shell morphology in endemic Greek land snails of the genus Codringtonia

Extensive variation in land snail shell morphology has been widely documented, although few studies have attempted to investigate the ecological and evolutionary drivers of this variation. Within a comparative phylogenetic framework, we investigated the temporal and spatial evolution of the shell morphology of the Greek endemic land snail genus Codringtonia. The contribution of both inter‐ and intraspecies shell differentiation in the overall shell variability is assessed. The effect of climate, space, and evolutionary history on the shell variability was inferred using a variance partitioning framework. For Codringtonia species, intraspecies divergence of shell traits contributes substantially to the overall shell variability. By decomposing this variability, it was shown that the overall shell size of Codringtonia clades is phylogenetically constrained, related to early speciation events, and strongly affected by large‐scale spatial variability (latitudinal gradient). The effect of climate on shell size cannot be disentangled from phylogeny and space. Shell and, to a larger extent, aperture shape are not phylogenetically constrained, and appear to be mostly related to conspecific populations divergence events. Shell shape is substantially explained by both climate and space that greatly overlap. Aperture shape is mainly interpreted by medium to small‐scale spatial variables. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 796–813.     

Evolutionary stasis,ecophenotypy and environmental controls on ammonite morphology in the Late Cretaceous (Maastrichtian) Western Interior Seaway,USA

We test for the presence of evolutionary stasis in a species of Late Cretaceous ammonoid cephalopod, Hoploscaphites nicolletii, from the North American Western Interior Seaway. A comprehensive dataset of morphological traits was compiled across the entire spatial and temporal range of this species. These were analysed in conjunction with sedimentologically and geochemically derived palaeoenvironmental conditions hypothesized to apply selective pressures. All changes in shell shape were observed to be ephemeral and reversable, that is, no unidirectional trend could be observed in any of the morphological traits analysed. Correlations between palaeoenvironmental conditions and morphological traits suggests ecophenotypic processes were at play; however, either environmental changes were too minor and/or provided no isolating mechanism to drive speciation. These data support mechanisms of stasis such as homogenizing gene flow or stabilizing selection under a fluctuating optimum (probably reflecting spatiotemporally heterogeneous palaeoenvironmental conditions). Finally, changes in shell size were not significantly associated with changes in shell-specific δ¹⁸O, despite a correlation between shell size and δ¹⁸O averaged across horizons. This suggests a mismatch in scales of geochemical sampling that supports caution when making broad interpretations based on averaged geochemical data.     

New insights into the diversity dynamics of Triassic conodonts

In this paper, we examine the diversity trends and the evolutionary patterns of Triassic conodonts through a newly powered large-scale data-set compiled directly from the primary literature. Paleodiversity dynamics analyses have been undertaken by working at the species level and using a system of time units based on biozone subdivisions for a fine temporal level resolution. The role of heterogeneous duration of taxa in diversity estimates has been evaluated through the probabilistic profiles. Results reveal three different stages in the diversity behaviour of Triassic conodonts from standing metrics delimited by two inflections at the mid-Anisian and mid-Carnian. Survivorship analysis supports this pattern. Origination–extinction metrics report a diversification pattern characterised by important fluctuations during the Lopingian–Induan (earliest Triassic), the early-middle Olenekian (Early Triassic) and the Anisian–Ladinian transitions (Middle Triassic), as well as in the early Late Triassic. In addition, two clear diversification peaks are observed in the late Carnian and in the end-Norian. Reported patterns are interpreted in the context of deep extinction and environmental instability by documenting the biological signal of the main diversification and turnover patterns observed from such records. This study emphasises the singularity behaviour of diversity trends derived from the conodont record.