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.     

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.     

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).     

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.     

Paedomorphosis of ammonoids as a result of sealevel fluctuations in the Late Devonian Wocklumeria Stufe

A remarkable diversification of several independent ammonoid lineages with high evolutionary rates occurred in the Late Devonian Wocklumeria Stufe. Many speciation events led to paedomorphic ammonoids that display a striking range of conch shapes, sculpture, and ornamentation. In the goniatite family Prionoceratidae, the transition from normal Mimimitoceras species to paedomorphic Balvia species provides an example of rapid size decrease combined with an early character developmental offset arising from progenesis. Adults of early Balvia species largely have the preadult ancestral morphology of Mimimitoceras , but later evolving species acquire distinct conch and ornamentation types. Progenetic ammonoid species also appeared within the clymeniid family Kosmoclymeniidae and probably in the Glatzielliidae. In the clymeniid family Parawocklumeriidae, evolution is characterized by the extension of tri-segmented and triangularly coiled whorls found only in juveniles of earlier species, to the adults of later species. This is interpreted as resulting from neoteny. The distribution of paedomorphic ammonoids in the Late Devonian Wocklumeria Stufe is closely correlated to relative sealevel changes. The regressive trend in the lower two-thirds of the Wocklumeria Stufe is interpreted as the cause of a diversification of the pelagic habitat during unstable conditions, and as an extrinsic factor inducing heterochronic change. Some ammonoids reacted by rapid maturation and faster reproductive rates, giving the opportunity to exploit a wider range of niches. The apparent consequence was the formation of several allopatric species. □ Ammonoidea, Late Devonian, evolution, heterochrony, sealevel changes.     

Buoyancy of some Palaeozoic ammonoids and their hydrostatic properties based on empirical 3D‐models

The interpretation of the function of the ammonoid phragmocone as a buoyancy device is now widely accepted among ammonoid researchers. During the 20^th century, several theoretical models were proposed for the role of the chambered shell (phragmocone); accordingly, the phragmocone had hydrostatic properties, which enabled it to attain neutral buoyancy, presuming it was partially filled with gas. With new three‐dimensional reconstructions of ammonoid shells, we are now able to test these hypothetical models using empirical volume data of actual ammonoid shells. We investigated three Palaeozoic ammonoids (Devonian and Carboniferous), namely Fidelites clariondi, Diallagites lenticulifer and Goniatites multiliratus, to reconstruct their hydrostatic properties, their syn vivo shell orientation and their buoyancy. According to our models, measurements and calculations, these specimens had aperture orientations of 19°, 64° and 125° during their lives. Although none of our results coincide with the aperture orientation of the living Nautilus, they do verify the predictions for shell orientations based on published theoretical models. Our calculations also show that the shorter the body chamber, the poorer was the hydrodynamic stability of the animal. This finding corroborates the results of theoretical models from the 1990s. With these results, which are based on actual specimens, we favour the rejection of hypotheses suggesting a purely benthonic mode of life of ammonoids. Additionally, it is now possible to assess hydrodynamic properties of the shells through ontogeny and phylogeny, leading to insights to validate theoretical modes of life and habitat through the animal's life.     

Cephalopod embryonic shells as a tool to reconstruct reproductive strategies in extinct taxa

An exhaustive study of existing data on the relationship between egg size and maximum size of embryonic shells in 42 species of extant cephalopods demonstrated that these values are approximately equal regardless of taxonomy and shell morphology. Egg size is also approximately equal to mantle length of hatchlings in 45 cephalopod species with rudimentary shells. Paired data on the size of the initial chamber versus embryonic shell in 235 species of Ammonoidea, 46 Bactritida, 13 Nautilida, 22 Orthocerida, 8 Tarphycerida, 4 Oncocerida, 1 Belemnoidea, 4 Sepiida and 1 Spirulida demonstrated that, although there is a positive relationship between these parameters in some taxa, initial chamber size cannot be used to predict egg size in extinct cephalopods; the size of the embryonic shell may be more appropriate for this task. The evolution of reproductive strategies in cephalopods in the geological past was marked by an increasing significance of small‐egged taxa, as is also seen in simultaneously evolving fish taxa.     

The Evolution and Development of Cephalopod Chambers and Their Shape

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.     

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.     

Analysis of a Carboniferous embryonic ammonoid assemblage implications for ammonoid embryology

An embryonic ammonoid assemblage was discovered in a carbonate concretion recovered from a dysoxic, relatively offshore marine shale of Virgilian (Upper Pennsylvanian) age in Kansas, USA. The assemblage consists primarily of two species of the Goniatitina, Aristocerassp. and Vidrioceras sp., whose initial chambers (protoconchs) differ in size and shape. Microscopic observations of serial thin sections of specimens at different growth stages reveal the sequence of embryonic shell development starting with the formation of the initial chamber and ending with the synchronous secretion of a prismatic proseptum and nacreous swelliig (primary varix) at the aperture. The mode of occurrence of the embryonic shells of the two species in the concretion suggests that these ammonoids produced numerous small offspring, a reproductive strategy similar to that in many extant coleoids. □ Ammonoids, embryonic shells, development, Carboniferous, Kansas.     

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 mode of life in ammonoids

The following structural features clearly indicate that ammonoid shells were adapted to withstand considerably higher hydrostatic pressures thanNautilus shells: (1) the corrugated and marginally fluted septa gave the shell wall efficient support against implosion; (2) the secondary connecting rings could grow a great deal in thickness; and (3) the last formed chambers remained full of liquid which supported the last septum. On the basis of the following characters it is concluded that ammonoids were incapable of swimming efficiently by jet-propulsion: (1) the retractor muscles were weakly developed; (2) the life position was unstable and highly variable; and (3) in animals with a ventral apertural rostrum the hyponome was probably absent. Ammonoids are considered here as having been pelagic cephalopods which lived in the upper 1000 m of the oceans, and which probably undertook considerable diurnal vertical migrations, similar to those inSpirula. Only some groups may have adopted a life in shallow epicontinental seas. In the late Mesozoic, ammonoids have been replaced by modern oceanic squids which are extremely numerous in the corresponding pelagic environment.     

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.     

Morphostructural constraints and phylogenetic overprint on sutural frilling in Late Jurassic ammonites

The functional significance of frilled septa and complex sutures in ammonoids has generated ongoing debate. The 'classic' hypothesis envisages ammonoid shells and septa as designed for resisting ambient hydrostatic pressure, complex sutures being the evidence of strength in shells for colonization of deep habitats. Here we address the 'suture problem', focusing on the analysis and interpretation of variables in our database of Late Jurassic ammonites not included in previous studies, such as whorl height ( W h ), whorl shape ( S ), shell coiling ( WD ), taxonomic grouping and basic planispiral shell shape. The results indicate that sutural complexity, as measured by the fractal dimension ( D f ) value of the suture line, is positively correlated with W h , and that the sutures of oceanic shells tend to provide, for a given W h value, lower D f estimates than do those of neritic shells. No general trend of increase in sutural complexity was noted for specimens recovered from swell areas belonging to oceanic fringes with respect to those that inhabited epicontinental shelves. In fact, Perisphinctoidea, the clade best represented in the database analysed, shows a higher D f mean value in neritic species than in epioceanic ones. Significant differences in sutural complexity were detected for groups of ammonites classified according to shell shapes ( WD , S ). Oxycones and discocones, streamlined potential swimmers, show the highest D f mean values, while spherocones and cadicones, which were presumably vertical vagrants, present the lowest ones. This indicates that sutural complexity was more related to shell geometry than to bathymetry.     

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.     

On the efficiency of the buoyancy apparatus in ammonoids: evidences from sublethal shell injuries

The five greatest sublethal injuries were selected from a collection of more than 12,000 predominantly Mesozoic injured or otherwise pathological ammonoids. The loss of shell mass from these survived injuries was calculated and compared with comparable tolerances in the recent Nautilus . These ammonoids tolerated a shell loss up to four times greater than in Nautilus . The maximum tolerated shell loss indicates an unexpected buoyancy compensation mechanism. The buoyancy of the selected specimens was calculated. The results show that the buoyancy of all the observed ammonoid shells was positive. In order to maintain neutral buoyancy after injury, these ammonoids had to fill the phragmocone with a volume of mass. Nautilus compensated a maximum mass loss requiring a liquid refill of 3% of the cameral capacity, the ammonoids compensated a maximum of observed mass loss requiring a liquid refill of more than 10% of cameral capacity. The ratio of chamber volume/siphuncular surface area in the ammonoid Lithacoceras is 0.043, indicating that the relative area of the siphuncular epithelium in Lithacoceras is significantly higher when compared with a ratio of 0.12-0.14 in the adult Nautilus . The phragmocone in ammonoids offered the ability of a much more active buoyancy regulation than in Nautilus .     

Thriving at high hydrostatic pressure: the example of ammonoids (extinct cephalopods)

Ammonoids are a group of extinct mollusks belonging to the same class of the living genus Nautilus (cephalopoda). In both Nautili and ammonoids, the (usually planospiral) shell is divided into chambers separated by septa that, during their lifetime, are filled with gas at atmospheric pressure. The intersection of septa with the external shell generates a curve called the suture line, which in living and most fossil Nautili is fairly uncomplicated. In contrast, suture lines of ancient ammonoids were gently curved and during the evolution of the group became highly complex, in some cases so extensively frilled as to be considered as fractal curves. Numerous theories have been put forward to explain the complexity of suture ammonoid lines. Calculations presented here lend support to the hypothesis that complex suture lines aided in counteracting the effect of the external water pressure. Additionally, it is suggested that complex suture lines diminished shell shrinkage caused by water pressure, and thus aided in improving buoyancy. Understanding the reason for complex sutures in ammonoids represents an important issue in paleobiology with potential applications to the problem of the resistance of hollow mechanical structures subjected to high pressure.     

Preferential predatory peeling: Ammonoid vs. nautiloid shells from the Upper Carboniferous of Texas,USA

Narrow groove-like excavations on ammonoid and coiled nautiloid shells are rare in Upper Carboniferous units from Texas, USA. The morphological characteristics of the excavation grooves typically are confined to the ventral and ventrolateral parts of the outer whorl of the shell, are narrower than the length, and have irregular edges where small segments or chips of shells have been removed. Analysis of these features reveals a statistically significant preferential occurrence on ammonoids (1.195% of ca. 3515 specimens) as compared to coiled nautiloids (0.506% of ca. 2965 specimens). The ammonoids typically have longer excavations that penetrate the phragmocone more frequently than those observed in the coiled nautiloids. The groove-like excavations were probably formed by the removal and peeling of shell material by one or more predatory or scavenging arthropods to obtain organic material (tissue and membranes) within the ammonoid and nautiloid body chambers and phragmocones. The excavations probably occurred when the cephalopod was alive (i.e., the cause of death) or shortly after the cephalopod's death. There is no evidence that the excavations are related to sheltering by the excavating organism.     

Post-hatching early life history of Cretaceous Ammonoidea

Post-hatching early life histories in Cretaceous Ammonoidea are discussed on the basis of density calculations of the shells in 71 species belonging to four separate suborders. The calculation was made under the assumption that a newly hatched ammonoid had a gas-filled chamber and a succeeding body-filled whorl terminating at the primary constriction. The results show that the density of the species examined at the hatching stage is almost constant and is relatively smaller than that of seawater, i.e. the animals are positively buoyant. This fact strongly suggests a planktic mode of life. In all species, the density increases gradually with growth and attains neutral buoyancy at 2.C2.5 mm in shell diameter. Thus, most ammonoids probably changed their mode of life from planktic to nektoplanktic or nektobenthic at this critical point. The rare occurrence of newly hatched specimens (ammonitellas) in many ammonoid assemblages may also support this interpretation. Planktic duration of a newly hatched ammonoid might be regulated by the animal's density at hatching, shell growth pattern, cameral volume (or hatching size), and rate of cameralliquid removal (or siphuncle diameter). The latter two seem to be very important factors in determining the biogeographical framework of species, as demonstrated in the Tetragonitaceae.□ Cretaceous, Ammonoidea, density calculation, early life history .