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.     

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.     

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.     

Some rare helical Ammonoidea from the lower Aptian (Lower Cretaceous) of the Paris Basin (NE France)

In the present paper, we describe and figure some rare lower Aptian (Lower Cretaceous) helical ammonoids from the Deshayesites deshayesi Zone of the Argiles à Plicatules Formation (Paris Basin, NE France). Those are the only helical ammonoids known in this zone so far. They were only occasionally evoked in literature and this is probably on the basis of the specimens here described that, according to us, the genus Toxoceratoides (Helicancylidae) was wrongly regarded as possessing an early helical spiral part. We demonstrate here that these specimens are not belonging to Toxoceratoides nor Helicancylidae: they are interpreted as ‘abnormally’ helical representatives of the genus Ancyloceras (Ancyloceratidae) or a new taxon of Ancyloceratidae.     

Ecological relations between hermit crabs and their shell-supplying gastropods: Constrained consumers

Hermit crabs are critically dependent upon gastropod shells for their survival and reproductive fitness. While anecdotal reports have suggested that hermit crabs may be capable of removing live gastropods from their shells to access the essential shell resource, no systematic experiments have been conducted to investigate this possibility. This paper reports experiments on both marine (Pagurus bernhardus) and terrestrial (Coenobita compressus) hermit crabs in which crabs were paired in the laboratory with the gastropods whose shells they inhabit in the field. Pairings included both shelled and naked crabs and spanned the full range of the gastropod life cycle. Neither marine nor terrestrial hermit crabs were successful at removing live gastropods from their shells. Furthermore, only a small fraction of the crabs (5.7%) were capable of accessing shells in which the gastropod had been killed in advance, with its body left intact inside the shell. Finally, although hermit crabs readily entered empty shells positioned on the surface, few crabs (14.3%) were able to access empty shells that were buried just centimeters beneath them. These results suggest that hermit crabs are constrained consumers, with the shells they seek only being accessible during a narrow time window, which begins following natural gastropod death and bodily decomposition and which typically ends when the gastropod's remnant shell has been buried by tidal forces. Further experiments are needed on more species of hermit crabs as well as fine-grained measurements of (i) the mechanical force required to pull a gastropod body from its shell and (ii) the maximum corresponding force that can be generated by different hermit crab species' chelipeds.     

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).     

A late devonian ammonoid faunula from himachal pradesh,india

From the uppermost parts of the Muth Quartzite Formation the following ammonoids are described which here are reponed from the Himalayas for the first time:Discoclymenia cucullata (v. Buch);Prionoceras (Imitoceras) intermedium Schindewolf;Pr. (I.) cf.rotundum Petter;Cyrtodymenia sp.; andDiscoclymenia striata (Münster). The levels containing ammonoids appear in the Famennian V(Clymenia zone). Their extension up to the earliest Tournaisian(Gattendorfia zone) is not likely although theoretically it cannot be excluded.     

Age,Growth and Spatial Distribution of the Life Stages of the Shortfin Mako,Isurus oxyrinchus (Rafinesque, 1810) Caught in the Western and Central Atlantic

The shortfin mako (Isurus oxyrinchus) is a highly migratory pelagic shark that preferentially inhabits oceanic regions in practically all oceans. The wide distribution range of this species renders it susceptible to coastal and oceanic fishing operations. The International Union for Conservation of Nature (IUCN) and the International Commission for the Conservation of Atlantic Tunas (ICCAT) consider this species to be highly vulnerable, especially due to its biological parameters, which are different from those of other sharks that occupy the same niche (e.g., Prionace glauca). Consequently, considerable declines in abundance have been detected over various parts of its range, most of which are linked to oceanic longline fishing. The species has conflicting life history parameters in studies conducted in the last 30 years, especially with regard to age and growth. The main discrepancies regard the interpretation of the periodicity of the deposition of band pairs (BPs) on vertebrae and the possibility of ontogenetic variations in growth. Shortfin mako sharks (n = 1325) were sampled by onboard observers of the Brazilian chartered pelagic longline fleet based in northeast Brazil from 2005 to 2011. Lengths were 79 to 250 and 73 to 296 cm (fork length, FL) for males and females, respectively, with a statistically significant difference in size between sexes and differences in the proportion of individuals in each size class. The onboard observers collected a subsample of vertebrae (n = 467), only 234 of which were suitable for analyses. Reliability between readings was satisfactory. However, it was not possible to validate periodicity in the formation of age bands in the sample. Thus, the von Bertalanffy growth function was used to calculate growth rates for the species through the interpretation of BPs in different scenarios: one BP per year (s1), two BPs per year (s2) and two BPs per year until five years of life (s3). Growth parameters varied for both females (Linf = 309.7[s3] to 441.6[s1]; k = 0.04[s1] to 0.13[s3]; t0 = -7.08[s1] and -3.27[s3]) and males (Linf = 291.5[s3] to 340.2[s1]; k = 0.04[s1] to 0.13[s3]; t0 = -7.08[s1] and -3.27[s3]). To advance the understanding of the use of habitat, the first analysis of the spatial distribution of the life stages of the shortfin mako sharks caught by commercial longline fishing operations in the South Atlantic was performed. The findings indicate that the portion of the population exploited by the fleets is predominantly juvenile and future actions should take the following issues into account: improvements in the efficiency of data collection procedures, the reestablishment of the onboard observer program, emergency investments in studies that can contribute to a better understanding of habitat use and life history theory.     

The ink sac clouds octopod evolutionary history

Difficulties in elucidating the evolutionary history of the octopods have arisen from problems in identifying informative morphological characters. Recent classifications have divided the largest group, the incirrate octopods, into five groups. These include the pelagic superfamily Argonautoidea and three gelatinous pelagic families (Vitreledonellidae, Bolitaenidae, Amphitretidae). All benthic incirrate octopods have been accommodated in the family Octopodidae, itself divided into four subfamilies, Octopodinae, Eledoninae, Bathypolypodinae and Graneledoninae, which are defined by the presence or absence of an ink sac, and uniserial or biserial sucker arrangements on the arms. We used relaxed clock models in a Bayesian framework and maximum likelihood methods to analyse three nuclear and four mitochondrial genes of representatives from each of the previous subfamilies. Strong evidence indicates that the family Octopodidae is paraphyletic and contains the gelatinous pelagic families. The subfamilies of Octopodidae recognised in earlier works do not reflect evolutionary history. The following clades were supported in all analyses: (1) Eledone/Aphrodoctopus, (2) Callistoctopus/Grimpella/Macroctopus/Scaeurgus, (3) Abdopus/Ameloctopus/Amphioctopus/Cistopus/Hapalochlaena/Octopus, (4) Enteroctopus/Muusoctopus/Vulcanoctopus, (5) Vitreledonella/Japetella, (6) Southern Ocean endemic and deep-sea taxa with uniserial suckers. These clades form the basis for a suite of taxa assigned family taxonomic rank: Amphitretidae, Bathypolypodidae, Eledonidae, Enteroctopodidae, Megaleledonidae and Octopodidae sensu nov. They are placed within the superfamily Octopodoidea.     

Chaos and stability of age-0 fish assemblages in a temperate deep reservoir: unpredictable success and stable habitat use

Large year-to-year variability in different fish species recruitment has been confirmed by previous studies while diurnal patterns of occupation in two basic reservoir habitats (pelagic and littoral) by different age-0 fish species in late summer are still unclear. Data collected over an 11-year period regarding late-summer age-0 fish assemblages in pelagic and littoral habitats of a reservoir were used to test the recruitment instability and to investigate diurnal habitat use. Trawling was conducted in the pelagic habitat at night while beach seining was conducted in the littoral habitat during day and night. Fluctuations in age-0 fish abundance and species composition were observed with both sampling methods; however, the following spatio-temporal patterns were relatively stable in most investigated years: (1) pelagic species (pikeperch; Sander lucioperca, small perch; Perca fluviatilis, bream; Abramis brama at night), (2) littoral species (large perch, asp; Leuciscus aspius, dace; Leuciscus leuciscus), (3) migratory species likely performing diel horizontal migrations (bleak; Alburnus alburnus), (4) species abundant in the littoral habitat both during day and night and also in pelagic habitat at night (roach; Rutilus rutilus) and (5) species detected in both habitats exclusively at night (ruffe; Gymnocephalus cernuus).     

Ammonoid biostratigraphy of the Middle Triassic Latemar platform (Dolomites, Italy) and its correlation with Nevada and Canada

An extensive study of the ammonoid fauna occurring in the lagoonal facies of a Middle Triassic isolated carbonate platform (Latemar platform, Dolomites, Italy) has been undertaken, and ammonoids from selected coeval successions (Punta Zonia, Marmolada, Rio Sacuz) have been illustrated. Ammonoids from Latemar have been collected in 20 distinct horizons (storm deposits) within the ca. 500 m thick lagoonal succession of the platform, thus providing a biostratigraphy of a series which is unusually expanded for this time interval, close to the Anisian-Ladinian boundary. Contrary to general opinion, some ammonoids of the Latemar and other coeval carbonate platforms of the Dolomites (Marmolada, Cernera), in particular Aplococeras avisianum, Lecanites misanii, Celtites spp., and Paranevadites sp., are also present in nearby basinal series. The same taxa have been found in North American localities deposited at the opposite margin of Panthalassa. The homotaxis of these ammonoids in North America and Latemar allow to establish a global scale correlation between the Southern Alps and North America with the highest resolution to date possible. In the context of this study, Aplococeras transiens n. sp. and Esinoceras nerinae n. sp are established.     

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.     

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.     

Ventral bite marks in Mesozoic ammonoids

Reports on the predators of ammonoids are rare, although ammonoids were abundant and diverse invertebrates in many Paleozoic and Mesozoic marine ecosystems. Most previous work on lethal ammonoid predation has focused on (sub)circular tooth marks which resulted from fish and mosasaur attacks. In the present study we discuss a relatively common type of bite mark in ammonoid shells, the ‘ventral bite mark’. This typically occurs in a restricted position on the ventral side of the outer body chamber whorl and does not affect either the aperture or the phragmocone. Ammonoid specimens revealing ventral bite marks used in this study were collected from a wide range of strata which range in age from the Lower Jurassic to the uppermost Cretaceous (close to the Cretaceous–Paleogene boundary). These ventral bite marks are absent in the Paleozoic collections studied. The vast majority of ventral bite marks are situated at the end of the body chamber, close to the phragmocone. This is interpreted as the result of predatory attacks on the back or blind side of ammonoids in their living position. The predators aimed for the vital parts and muscle attachments to obtain the edible soft tissues. The agents for most of the ventral bite marks to ammonoids are probably coleoid cephalopods (especially teuthoids) and predatory fishes to a lesser extent.     

The lipid compositions of high-Antarctic notothenioid fish species with different life strategies

Lipid and fatty acid compositions of five notothenioid fishes from the Antarctic Weddell and Lazarev Seas were investigated in detail with regard to their different modes of life. The pelagic Aethotaxis mitopteryx was the lipid-richest species (mean of 61.4% of dry mass, DM) followed by Pleuragramma antarcticum (37.7%DM). The benthopelagic Trematomus lepidorhinus had an intermediate lipid content of 23.2%DM. The benthic Bathydraco marri (20.8%DM) and Dolloidraco longedorsalis (14.5%DM) belonged to the lipid-poorer species. Triacylglycerols were the major lipid class in all species. Important fatty acids were 16:0, 16:1(n-7), 18:1(n-9), 18:1(n-7), 20:5(n-3) and 22:6(n-3). The enhanced proportions of the long-chain monounsaturated fatty acids, 20:1 and 22:1, in the lipid-rich pelagic fishes clearly reflected the ingestion of the two copepod species, Calanoides acutus and Calanus propinquus, which are the only known Antarctic zooplankters rich in these fatty acids. Although wax esters are the major storage lipid in many prey species, they were absent in all notothenioid fishes studied. Thus, wax esters ingested with prey are probably converted to triacylglycerols via fatty acids or metabolised by the fishes. The enhanced lipid accumulation with increasingly pelagic lifestyle has energetic advantages, especially with regard to improved buoyancy. It is still unknown to what extent these lipids are utilised as energy reserves, since it has been suggested that not only the benthic but also the pelagic Antarctic fishes are rather sluggish, with a low scope for activity and hence low metabolic requirements. Accepted: 14 May 2000     

Evidence of predation in Early Cretaceous unionoid bivalves from freshwater sediments in the Cameros Basin,Spain

Here, we present evidence of possible vertebrate predation on freshwater bivalves from the Lower Cretaceous strata of the Cameros Basin (Spain). The described collection contains the largest number of vertebrate‐inflicted shell injuries in freshwater bivalve shells yet reported in the Mesozoic continental record. Several types of shell damage on fossil shells of Protopleurobema numantina (Bivalvia: Unionoida) are described and their respective modes of formation interpreted in the context of morphological attributes of the shell injuries and the inferred tooth morphology of predators that could have inflicted such injuries. Detailed study of these bite marks shows similarities with the well‐documented injuries in the shells of marine molluscs, namely ammonoids, that have likewise been attributed to reptilian predators. The most parsimonious interpretation suggests crocodiles as the vertebrates interacting with the bivalves in the Cameros Basin. □Barremian–Aptian; bite marks; freshwater bivalves; predation; reptile; Unionoida.     

Evolution of pelagic direct development in the starfish Pteraster tesselatus (Asteroidea: Velatida)

Despite a diversity of larval forms, remarkably conservative features of asteroid development define a larval body plan that occurs throughout the class. However, recent work on the starfish Pteraster tesselatus has documented a highly derived pattern of development. Several features, including radial symmetry, parallel embryonic and adult axes of symmetry, absence of a preoral lobe, and formation of coeloms in the adult orientation from seven separate enterocoels, have not been reported in asteroids before. The complete absence of the larval body plan features that are found in other asteroids, indicates that P. tesselatus develops directly from the embryo to the juvenile and has a pelagic, nonfeeding (lecithotrophic), but nonlarval mode of development. I postulate that direct development evolved over an extended period in a lineage of brooding, deep-sea velatid (probably pterastcrid) ancestors of P. tesselatus. Selection for increased developmental efficiency (loss of nonfunctional larval features) in the brooded offspring, could explain the lack of larval settlement structures, the nonlarval arrangement of coeloms, the lack of a preoral lobe, the transverse orientation of the juvenile disc, and the lack of bilateral symmetry. The pattern of coclomogenesis could have been derived from that of other velatids (e.g. solasterids) by relatively simple changes in timing and orientation of entcroeoel formation. Rotation and posterior translation of the coelomic fate map of the archenteron prior to enlerocoel formation would produce the coelomic compartments in the adult orientation that characterizes direct development in P. tesselatus. These unusual developmental features lead to a radically different interpretation for the evolution of the pelagic ‘larva’ of P. tesselatus: (1) evolution of benthie brooding, (2) extreme simplification of development involving the loss of all larval features from the life cycle, and (3) subsequent re-evolution of pelagic development. In the case of P. tesselatus, where all larval structures were lost, there do not seem to be functional constraints preventing the re-evolution of pelagic development. Analysis of pelagic and benthie larvae, in other asteroids, suggests that major ecological transitions in life histories need not be associated with substantia] changes in morphology. The loss of pelagic development should have occurred repeatedly and should be readily reversible. These findings have interesting implications for the loss and evolution of pelagic dispersal in the life histories of marine benthie invertebrates.     

A forced association between the slippersnail Crepidula convexa and the hermit crab Pagurus longicarpus?—possible influence from a third party

The suspension-feeding slippersnail Crepidula convexa is commonly associated with hermit crabs (Pagurus longicarpus) living in periwinkle shells (Littorina littorea) at our study site in Nahant, MA, USA. In 15 field surveys conducted at Nahant in 2000, 2001 and 2003, we found that (1) more than 61.8% of individuals of C. convexa resided on shells occupied by hermit crabs, as opposed to the shells of live periwinkles, empty periwinkle shells or other solid substrates; (2) an average of 8.3% of hermit crabs carried at least one individual of C. convexa; and (3) 39.1-75.0% of hermit crabs carrying C. convexa were carrying “large” individuals (snails with wet weight >10% of the weight of the periwinkle shells they occupied). However, it is unlikely that individuals of C. convexa seek out shells occupied by hermit crabs to colonize, and they showed no preference for empty periwinkle shells over other solid substrates in the laboratory. Moreover, in the laboratory the hermit crabs preferentially occupied intact shells bearing individuals of C. convexa only when the alternatives were shells that had been drilled by naticid snails. Thus, neither party preferentially associates with the other: rather, extensive predation by naticid snails on periwinkles at Nahant appears to limit the availability of suitable shells for the hermit crabs, forcing them to inhabit shells bearing “large” individuals of C. convexa. Individuals of C. convexa may benefit from this inadvertent association with hermit crabs: by facilitating snail dispersal, transport by hermit crabs should reduce the potential for inbreeding, an important consideration for a species that lacks free-living larvae in its life history.     

Spirula—a window to the embryonic development of ammonoids? Morphological and molecular indications for a palaeontological hypothesis

Nautilus is not suitable as a model organism to infer biological functions, embryonic development, or mode of life in ammonoids. A brief review of the available morphological data is given and molecular data are added to discuss the usefulness of Spirula as a biological proxy for ammonoids. Indeed, there are many morphological hints indicating that Spirula could be a useful model organism for approaching the embryonic development of ammonoids. The molecular data seem to support this hypothesis. However, a universal model character of Spirula cannot be detected as, e.g., the mode of feeding probably differs between Spirula and ammonoids.