Covariation of morphological characters in the Triassic ammonoid Czekanowskites rieberi

The Triassic ammonoid Czekanowskites rieberi displays a covariation of morphological charac ters, which is rather common in ammonoids. Its morphological spectrum ranges from laterally compressed, involute, weakly ribbed forms to depressed, semiinvolute, strongly ribbed forms. In order to study this covariation, fifteen axially cut specimens have been analyzed by means of image analysis, which allows us to obtain the ontogenetic record of radii, area and perimeter of the individual whorl cross-sections. A logarithmic model of growth has been applied. Our data indicate that, owing to the covariation, the radii from the origin to the venter and to the umbil ical seam of a given whorl section vary inversely in order to maintain the relative position of the center of gravity of the whorl cross-section both throughout the ontogeny of single specimens and within the population. This influences hydrostatic parameters, such as the position of the center of mass and the orientation and stability of the shell. Since the ontogenetic record of the angular length of the body chamber is not known, we have calculated those hydrostatic varia bles using two mutually exclusive assumptions: (1) the angular length of the body chamber was constant throughout ontogeny and (2) the volume of the body chamber grew monotonically with the revolution angle. Fluctuations of the three hydrostatic variables were always less important in the first assumption. In any case, the spectrum of, for example, theoretical orien tations is comparable to those observed in the species of present-day Nautilus. The range of adult body-chamber length observed in C. rieberi is much narrower than the theoretical adult body-chamber length calculated under the second assumption which indicates that a certain control over this parameter existed in the natural population, probably in order to maintain a narrow range in orientation and stability. The excess or deficit in soft-body weight was probably compensated by inverse variations in shell-wall weight. The main conclusion is that, despite the extreme morphological variability, hydrostatic and, possibly, hydrodynamic properties of the population remained within narrow limits.     

Morphogenetic significance of the conchal furrow in nautiloids: evidence from early embryonic shell development of Jurassic Nautilida

As reported by many workers over the past two centuries, the inner part of the shell of various straight and coiled Palaeozoic to tertiary nautiloid taxa bears a continuous mid-ventral furrow that extends into the phragmocone and the body chamber nearly to the aperture. Study of the early embryonic shell development of Jurassic Nautilida shows that the most apical part of this so-called conchal furrow originates from the inner part of the initial, calcified shell apex, in line with the inner ventral termination of the central linear depression of the cicatrix, the initial site of shell deposition. The conchal furrow corresponds to a morphological feature arising as a developmental by-product. Rare specimens of scattered ammonoid species (and possibly of bactritoids) display a similar feature, whereas their protoconch lacks a cicatrix. However, the protoconch of recent cuttlefish, Sepia officinalis, often displays a longitudinal fold of the primary shell epithelium. A longitudinal groove or a pair of grooves appears connected with this cicatrix-like structure. Although the mid-ventral ridge in ammonoids must probably be viewed as an incidental 'fabricational noise', whether or not it originates from a so far undocumented optional ridge on the protoconch or from some other structure related to shell development remains an open question.     

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

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

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.     

Post-pathological keel-loss compensation in ammonoid growth

Among the various pathologies documented in ammonoids, impairs affecting the apertural margin may have long-lasting sequelae on subsequent shell geometry. An interesting healing pattern, known as sculptural compensation, led to the permanent replacement of an ornament by adjacent sculptural elements. Moreover, in several ventrally impaired individuals the symmetry was preserved. Those developed annular ribs in place of any previous ventral ornamentation (keel, sulcus or smooth area). This phenomenon is known from diverse ammonite families. Monestieria resouchei (Monestier 1931), type species of 'Monestieriinae' Sapunov 1965, displays exactly that type of annularly-ribbed morphology and has been shown to be otherwise similar to species of Grammoceratinae Buckman 1904 occurring in the same beds, thus corroborating its pathological nature and leading to the rejection of that taxon. Now, keel absence in Praehaploceras Monestier 1931 and Buckmanites Guex 1973 cannot be explained by the same process as they do not have annular ribs. Moreover, the absence of any clue of malformation, their relative frequency and specific characteristics exclude the previously suggested synonymies with Pseudolioceras Buckman 1889 as equivalent pathological forms. In consequence, their rehabilitation is herein proposed. They should be included within Harpoceratinae Neumayr 1875.     

How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

Late Palaeozoic mollusc reproduction: cephalopod egg-laying behavior and gastropod larval palaeobiology

Faunal analysis of an oxygen‐depleted marine Lower Carboniferous succession (Late Mississippian Ruddle Shale) suggests how some cephalopod taxa laid their eggs during the Late Palaeozoic. At the Ruddle Shale collecting site in Arkansas, USA, the facies and overall fauna suggest severe oxygen depletion at the sediment/water interface. The Ammonoidea, with their small egg size, were probably laid in suspended gelatinous egg‐filled masses in the water column above the bottom or by attachment of the egg masses to floating debris. The ammonitella embryos developed within the suspended or attached egg mass; hatched individuals became part of the free‐swimming plankton biota. Based on shell morphology the Bactritoidea probably followed the same reproductive pattern. Coiled nautiloids (the Nautilida) and most orthoconic nautiloids (mostly the Pseudorthocerida) probably did not lay their eggs in the mid water column or as floatant attachments. This conclusion is based on the fact that, with one exception, all shells recovered of these two nautiloid orders are well past hatching. Gastropods in the Ruddle Shale are very small and cannot be visually detected in the field. However, microgastropods are abundant in washed residues. Most specimens are much smaller than 1 mm. The largest caenogastropod specimen is 1.3 mm high. These caenogastropods represent isolated larval shells and a successful metamorphosis was impossible because of oxygen depletion on the bottom. Allegations that a size of more than 1 mm is too large for pelagic larvae are refuted by examples of planktotrophic larval shells of modern gastropods (more than 1 mm high) and Triassic caenogastropods (up to 2 mm high) from the Cassian Formation (Northern Italy, South Alps). Repository information is given for the type‐material of the gastropod species Nuetzelina striata  Bandel, 2002 and Anozyga arkansasensis  Bandel, 2002 which were both erected based on specimens from the Ruddle Shale that were illustrated by Nützel & Mapes in 2001.     

Tithonian ammonoid biostratigraphy in eastern Himalayan Tibet

The Tithonian and Lower Berriasian sediments in the eastern Himalayas of Tibet contain an extensive sequence of ammonoid fauna. New collections in situ through the Lanongla, Pure, Gucuo and Jiapeila sections have facilitated a major revision of the ammonoid assemblages. Probably due to depositional facies segregation, the Belemnopsis galoi-bearing beds can be regarded as the oldest Tithonian sediments in which the basal Tithonian ammonoid Kossmatia is not present. The Lower Tithonian includes the Virgatosphinctes-Aulacosphinctoides and Uhligites-Aulacosphinctes assemblages; the Upper Tithonian includes the Haplophylloceras pingue, Blanfordiceras wallichi and Haplophylloceras strigile-Corongoceras-Himalayites assemblages. The Spiticeras assemblage is suggested to be from the Lowermost Berriasian. The new ammonoid assemblages at well-defined levels in Himalayan Tibet provide some crucial links for correlation with other regions of the SW Pacific domain where these ammonoid genera have been widely distributed.     

Biostratigraphy and important biotic events in the Western Verkhoyansk Region around the Sakmarian–Artinskian boundary

A diverse marine invertebrate fauna was found in the Echij Formation (Sakmarian–Artinskian) at the Arkachan, Chelge, and Nizhnyaya Dielendzha sections, all Western Verkhoyansk Region, North-East of Russia. The biostratigraphic sequence of ammonoid, brachiopod, bivalve, and foraminiferal assemblages in the Echij Formation of the Western Verkhoyansk Region is studied. Five ammonoids units are identified in the Echian Regional Stage (“Horizon”): Uraloceras subsimense, Uraloceras omolonense, Neoshumardites triceps hyperboreus, Eotumaroceras endybalense, and Eotumaroceras subyakutorum beds. The first two divisions contain ammonoids of the Arkachanian association, and the last three contain the Endybalian association. The boundary between the Sakmarian and Artinskian stages is established at the base of Neoshumardites triceps hyperboreus beds. In the Sakmarian interval of the Echij Regional Stage of the Verkhoyansk Region, a brachiopod biostratigraphic sequence similar to that of the Kolyma–Omolon Region is observed: Jakutoproductus insignis, Jakutoproductus terechovi, and Jakutoproductus rugosus zones. In the lower part of the Artinskian stage, Uraloproductus stuckenbergianus beds are identified, which are characterized by a rich brachiopod assemblage (Peregoedov et al., 2009), not typical for Verkhoyansk Region. The bivalves, identified in the Lower Echij Subformation, presumably belong to the Merismopteria permiana, Cypricardinia eopermica, and Cypricardinia borealica zones of the Ogonerian Horizon of the Kolyma–Omolon Region. Bivalves of the Middle and Upper subformations are characteristic of the Edmondia gigantea and Aphanaia lima zones of the Koargychanian Regional Stage (“Horizon”). The foraminifera complex identified in the Echij Formation is compared with the complex of the lower part of the Sandy Foraminifera horizon of the northeast of the Siberian platform, to which the Tustakh Formation belongs. The beginning of the wide distribution of the Early Permian foraminifera in the Western Verkhoyansk Region was recorded at the base of the Artinskian. The Sakmarian–Artinskian boundary interval in the Verkhoyansk Region is characterized by three significant biotic events: the replacement of the brachiopod Jakutoproductus by Inoceramus-like bivalves, the first appearance of the Endybalian ammonoid association, and biotic invasions from the Uralian and North American regions. An important factor of the Late Sakmarian–Early Artinskian events was a large sea level rise (the Echian transgression), which significantly changed the environmental conditions for East Siberian marine invertebrates, and contributed to the spread of new faunas.     

Experimental fragmentation patterns of modern Nautilus shells and the implications for fossil cephalopod taphonomy

Shells of modern Nautilus pompilius from the Philippines were experimentally fragmented designed to mimic: (1) transport with sediment; (2) sediment loading; and (3) collision during floating. The breaking patterns by other mechanisms (predation and implosion by hydrostatic pressure) documented in the literature were also considered. The breaking patterns produced by various mechanisms are distinct and can therefore be differentiated. The results allow identification of the distinct mechanisms responsible for specific fragmentation not only in modern Nautilus but also in fossil cephalopods. This is a new approach for the more complete recognition of post-mortem transport of fossil cephalopods and their early taphonomic history, and contributes to our understanding of their palaeobiogeography and palaeoecology.