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.     

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.     

Quantification of ontogenetic allometry in ammonoids

Ammonoids are well‐known objects used for studies on ontogeny and phylogeny, but a quantification of ontogenetic change has not yet been carried out. Their planispirally coiled conchs allow for a study of “longitudinal” ontogenetic data, that is data of ontogenetic trajectories that can be obtained from a single specimen. Therefore, they provide a good model for ontogenetic studies of geometry in other shelled organisms. Using modifications of three cardinal conch dimensions, computer simulations can model artificial conchs. The trajectories of ontogenetic allometry of these simulations can be analyzed in great detail in a theoretical morphospace. A method for the classification of conch ontogeny and quantification of the degree of allometry is proposed. Using high‐precision cross‐sections, the allometric conch growth of real ammonoids can be documented and compared. The members of the Ammonoidea show a wide variety of allometric growth, ranging from near isometry to monophasic, biphasic, or polyphasic allometry. Selected examples of Palaeozoic and Mesozoic ammonoids are shown with respect to their degree of change during ontogeny of the conch.     

Soft‐tissue imprints in fossil and Recent cephalopod septa and septum formation

Several soft‐tissue imprints and attachment sites have been discovered on the inside of the shell wall and on the apertural side of the septum of various fossil and Recent ectocochleate cephalopods. In addition to the scars of the cephalic retractors, steinkerns of the body chambers of bactritoids and some ammonoids from the Moroccan and the German Emsian (Early Devonian) display various kinds of striations; some of these striations are restricted to the mural part of the septum, some start at the suture and terminate at the anterior limit of the annular elevation. Several of these features were also discovered in specimens of Mesozoic and Recent nautilids. These structures are here interpreted as imprints of muscle fibre bundles of the posterior and especially the septal mantle, blood vessels as well as the septal furrow. Most of these structures were not found in ammonoids younger than Middle Devonian. We suggest that newly formed, not yet mineralized (or only slightly), septa were more tightly stayed between the more numerous lobes and saddles in more strongly folded septa of more derived ammonoids and that the higher tension in these septa did not permit soft‐parts to leave imprints on the organic preseptum. It is conceivable that this permitted more derived ammonoids to replace the chamber liquid faster by gas and consequently, new chambers could be used earlier than in other ectocochleate cephalopods, perhaps this process began even prior to mineralization. This would have allowed faster growth rates in derived ammonoids.     

The ammonitella: a model of formation with the aid of the embryonic shell of archaeogastropods

Recent archaeogastropoda secrete their first conch (primary shell) without producing increments of growth. This conch remains attached to the epithelium responsible for its production until completed. Only afterwards does tissue detach from the edge of the shell and the conch begins to function as a protective exoskeleton. After a potential swimming phase by the veliger larva, the organic primary shell is deformed by mechanical means through muscular tension from the inside and the outside of the conch. It then becomes mineralized by aragonitic crystallites and, thus, functional for the use of a benthic animal. The embryonic conch of ammonites (ammonitella) is devoid of increments of growth. The inner lip (dorsal side) of the aperture became flattened after the evenly rounded primary conch had been secreted. The primary organic shell was mineralized by aragonitic crystallites from within. All these features of the formation of the ammonitella can be interpreted in the light of early shell formation of recent archaeogastropods.     

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.     

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.     

The Micro/Nanostructure Characteristics and the Mechanical Properties of Hemifusus tuba Conch Shell

<正> The mollusk shell mobilizes calcium from environment for skeletal mineralization.This occurs through synthesizing solidsin solution in the presence of organic molecules of specific interior regions of the conch shell.The ultrastructure and microhardnessof the Hemifusus tuba conch shell living in the Huang/Bo sea area are investigated in the paper.It is shown that thecomposition and microstructure of the mollusk shell vary in different positions.The prodissoconch shell consists only of aragonitewith the crossed-lamellar microstructure.While the spiral shell and the body shell of the Hemifusus tuba conch shell arecomposed of one calcite layer and several aragonite layers.The calcite layer consists of cylindrical grains,but the aragonitelayers are crossed-lamellar ultrastructure at three size scales.The minimum structure size (the third-order lamella) is at about20 nm - 80 nm.The margin of shell aperture is only composed of calcite with cylindrical grains.This natural optimization of theshell microstructure is intimately due to the growth of the Organic matrix.At different positions the microhardness of molluscshell is different due to different crystal structures and crystal arrangements.The growth process of shells allows a constantrenewal of the material,thus enabling their functional adaptation to external environments.     

Description of the early ontogenic part of the Tentaculitids, with implications for classification

Ontogenic development and classification of tentaculitids at high systematic levels are reevaluated in the light of new findings on shell structure and morphology of larval parts, and these features are here regarded as being of primary importance for taxonomy. Class Tentaculita Bouček, 1964 is subdivided into two subclasses, of which subclass Chionioconarida Farsan, 1994 is distinguished by a tubular larval process closed at the apex and covered with microrings. The process is differentiated into a prolarval, metalarval and epilarval part, of which the latter coincides with metamorphosis. Morphology of the larval parts suggests that metamorphosis proceeds in two different manners, giving rise to superorders within this subclass. Within superorder Trompetoconarida Farsan, 1994 a bilaterally symmetrical larval cone develops with an aperture oblique to the long axis of the conch; following metamorphosis the conch becomes radially symmetrical and the aperture perpendicular to the axis; secondary shell, septa and pseudopunctae develop in the adult phase, and the structure of the shell is lamellar. In contrast, within the second superorder, Lirioconarida Farsan, 1994 the epilarval tube develops into a larval bulb with no changes in symmetry and position of the aperture; secondary shell, septa and pseudopuncta are absent. The microstructure of the shell is lamellar in the larval part whereas in postlarval parts it is either sigmoidal or lamellar. The subclass Dacryoconarida Fisher, 1962 possesses a subspherical, tear- or drop-like embryonic chamber which may have a caudal process. The microstructure of the embryonic chamber is variable within this group, being lamellar in some taxa whereas in others, a single layer of shell is present. The postembryonic parts of the lamellar forms possess nacreous or sigmoidal structures.     

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.     

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 .     

陕西洛南下寒武统辛集组疑难管状化石桶螺(Cupitheca)

本文对采自陕西洛南上湾剖面下寒武统辛集组疑难管状化石Cupitheca进行了研究,系统厘定和描述了两个种,分别为Cupitheca holocyclata和Cupitheca costellata,并对Cupithecidae科进行了重新修订。本文所描述标本的壳体多为次生磷酸盐化保存,壳体表面纹饰保存较好;C.holocyclata管体表面为横向纹饰,C.costellata管体表面为纵向纹饰。C.holocyclata全球广布,如南澳大利亚、南极乔治王岛、格陵兰东北部、加拿大纽芬兰等地皆有报道,产出于下寒武统中上部地层,具有一定的洲际地层对比意义。本文为C.holocyclata在中国首次发现。C.costellata仅发现于华北,即本文所研究的洛南辛集组及其同期地层安徽淮南与霍邱的雨台山组。     

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.     

Predator-induced behavioral and morphological plasticity in the tropical marine Gastropod Strombus gigas

Florida queen conch stocks once supported a significant fishery, but overfishing prompted the state of Florida to institute a harvest moratorium in 1985. Despite the closure of the fishery, the queen conch population has been slow to recover. One method used in the efforts to restore the Florida conch population has been to release hatchery-reared juvenile conch into the wild; however, suboptimal predator avoidance responses and lighter shell weights relative to their wild counterparts have been implicated in the high mortality rates of released hatchery juveniles. We conducted a series of experiments in which hatchery-reared juvenile conch were exposed to a predator, the spiny lobster (Panulirus argus), to determine whether they could develop behavioral and morphological characteristics that would improve survival. Experiments were conducted in tanks with a calcareous sand substrate to simulate a natural environment. Conditioned conch were exposed to caged lobsters while conch in the control tanks were exposed to empty cages. Conditioned conch moved significantly less and buried themselves more frequently than the naive control conch. Morphometric data indicated that the conditioned conch grew at a significantly slower rate than the naive conch, but the shell weights of the two groups were not significantly different. This implies that the conditioned conch had thicker or denser shells than the control group. As a result, the conditioned conch had significantly higher survival than naive conch in a subsequent predation experiment in which a lobster was allowed to roam free in each tank for 24 hours. In the future, the conditioning protocols documented in this study will be used to increase the survival of hatchery-reared conch in the wild.     

Nabelkanten-präferenz der formaverticata hölder 1956 bei dactylioceraten (ammonoidea,toarcien)

The favour position of the formaverticata Hölder 1956 on the umbilical edge at the ammonite genusDactylioceras (Upper Liassic, Franconia) is reduced on a punctual restriction of the peristomal mantle-epithelium. It seems to be a fact of an atteck of parasites, because

1. the anomaly is starting on one of the most secure positions of the conch, mostly without a visible infraction of the shell,
2. the abundance of these anomalies is very different at single localities.

     

The genus Luppovia and the phylogeny of Cretaceous heteromorphic ammonoids

The unique preservation of heteromorphic shells of Luppovia Kakabadze, Bogdanova & Mikhailova 1978 from the Aptian deposits of Bolshoj Balkhan (western Turkmenia) made it possible to study the microstructure, internal shell characteristics and suture ontogeny of this genus. Microstructural investigations revealed that the dorsal shell wall in Luppovia is composed of three layers in contrast to that of monomorphic ammonoids, which is single-layered. This ability of secreting different numbers of layers in monomorphs and heteromorphs demonstrates plasticity of secreting activity of the mantle epithelium. Data obtained from the study of internal shell structure and suture ontogeny have been compared. Both these methods of investigation lead to the conclusion that the genus Luppovia belongs to the order Ammonitida. The results obtained confirm the viewpoint that the Cretaceous heteromorphic superfamilies Turrilitaceae and Ancylocerataceae have independent origins and belong to two different orders, Lytoceratida and Ammonitida, respectively. □ Ammonoidea , Luppovia, shell structure, ontogeny, phylogeny, Cretaceous.     

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.      

Irregular calcareous sheets preserved in a conch of the Cretaceous ammonoid Hypophylloceras from Hokkaido,Japan

The mantle tissue is essential for understanding the diverse ecology and shell morphology of ammonoid cephalopods. Here, we report on irregular calcareous sheets in a well-preserved shell of a Late Cretaceous phylloceratid ammonoid Hypophylloceras subramosum from Hokkaido, Japan, and their significance for repairing the conch through the mantle inside the body chamber. The sheets are composed of nacreous layers arranged parallel to the irregularly distorted outer whorl surface. The nacreous sheets formed earlier are unevenly distributed and attached to the outer shell wall locally, whereas the last formed sheet covers a wide area of the outer shell wall. The absence of any interruption of ribbing around the irregular area suggests that these sheets were secreted inside the body chamber from the inner mantle. Gross morphological and X-ray computed tomography observations revealed that the spacing of septal formation was not affected by this event. The complex structure of the irregular sheets suggests a highly flexible mantle inside the body chamber.     

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.