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![Belemnoid fossils from the late Triassic Konservat-Lagerstätte Polzberg near Lunz am See. (A) Hook morphology in phragmoteuthid arm crowns. (B) Semi in situ preserved arm crown. Each arm comprises two rows of hook armory. Terms adapted from [30]. (C) Belemnoid fossil remains, assigned to the taxon Phragmoteuthis bisinuata with its characteristic fan-shaped three-parted proostracum. White arrows pointing to position of carbonized structure. https://doi.org/10.1371/journal.pone.0264595.g002](profile/Petra-Lukeneder/publication/360079692/figure/fig2/AS:1147152717877249@1650513929206/Belemnoid-fossils-from-the-late-Triassic-Konservat-Lagerstaette-Polzberg-near-Lunz-am-See.png)
Belemnoid fossils from the late Triassic Konservat-Lagerstätte Polzberg near Lunz am See. (A) Hook morphology in phragmoteuthid arm crowns. (B) Semi in situ preserved arm crown. Each arm comprises two rows of hook armory. Terms adapted from [30]. (C) Belemnoid fossil remains, assigned to the taxon Phragmoteuthis bisinuata with its characteristic fan-shaped three-parted proostracum. White arrows pointing to position of carbonized structure. https://doi.org/10.1371/journal.pone.0264595.g002
Source publication
Although hyaline cartilage is widely distributed in various invertebrate groups such as sabel-lid polychaetes, molluscs (cephalopods, gastropods) and a chelicerate arthropod group (horseshoe crabs), the enigmatic relationship and distribution of cartilage in taxonomic groups remains to be explained. It can be interpreted as a convergent trait in an...
Contexts in source publication
Context 1
... IRIS system (Interaktives Rohstoffinformations System) hints to an early historical adit (No. 071/3008a) for the production of black coal, active in the first half of the 19 th century, which was confirmed by the literature [28]. The outcrop Polzberg locality (Fig 2A) was first mentioned by [28]. Further historical adits for the recovery of fossil specimens were dug in 1885 (Geological Survey of Austria, GBA) and in 1909 (Natural History Museum Vienna, NHMW) by Joseph Haberfelner [25] and references therein at approx. ...
Context 2
... specimens stem from the Carnian Reingraben shales at the Polzberg locality (= Polzberggraben; historical and new findings). Bed-by-bed collected specimens found by the authors from distinct levels: Po -50-0 cm, Po 60-80 cm, Po 80-100 cm, Po 100-120 cm, Po 140-160 cm (Fig 2). Seven fossils come from Cave del Predil (northern Italy). ...
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Citations
... Discovered 150-years ago, the Polzberg locality ( Fig. 1) has yielded many belemnoid hard tissues including phragmocones, proostraca, cephalic cartilages and arm hooks (Doguzhaeva & Summesberger, 2012;Doguzhaeva et al., 2006Doguzhaeva et al., , 2007bLukeneder & Lukeneder, 2022b;Mojsisovics von Mojsvar, 1882;Suess, 1865). Soft tissues from ammonoids and belemnoids have also (Doguzhaeva et al., 2007a;Lukeneder & Lukeneder, 2022a). ...
... The material derives from the Reingraben shales of the late Triassic (Julian II, Carnian) Konservat-Lagerstätte Polzberg ( Fig. 1; GPS N 47° 53′ 5.90″ and E 15° 4′ 27.70″) (Lukeneder & Lukeneder, 2021, 2022a, 2022bLukeneder et al., 2020) at 712 m above sea level, which was first discovered by Stur (1874). The site is located about 4.5 km northeast of Lunz am See within the Polzberg Basin. ...
... Hence, the perspective to find a hook morphotype unambiguously characterising phragmoteuthids is attenuated Fuchs et al., 2021). (2) The various cephalic cartilages described from the Alpine Triassic (Lukeneder & Lukeneder, 2022b, 2022cRieber, 1970) are mostly preserved without associated shell remains which makes it difficult to unambiguously identify a morphotype that belonged to phragmoteuthids. (3) Until now, rostrum-less specimens from the Alpine Triassic were tentatively assigned to the genus Phragmoteuthis Mojsisovics (1882). ...
Coleoid cephalopods are widespread from the Mesozoic till today. The extinct group of the Phragmoteuthida is thought to represent either stem-neocoleoids, stem-decabrachians, or stem-octobrachians. The well-known, almost complete specimens of Phragmoteuthis bisinuata from the Carnian Polzberg Konservat-Lagerstätte near Lunz am See (Lower Austria, Northern Calcareous Alps) and Cave del Predil (Northern Italy, Julian Alps) come from historical collections. These specimens do not reflect the entire coleoid assemblage within this environment. In order to obtain a more complete picture of the Carnian coleoid fauna, 430 coleoid specimens from the Polzberg locality and 60 specimens from contemporaneous localities around Cave del Predil were studied in detail. All available elements (phragmocones, proostraca, cartilages, hooks, beaks, ink sacs) attributed to the coleoid fauna were recorded, measured and evaluated taxonomically and taphonomically. Reviews of historical collection material permitted comparisons with recently collected material. The notation of co-occurrences of other faunal elements yielded insights into the palaeoecological context of this Upper Triassic environment within the Polzberg Basin. The new material from Polzberg does not support the previous assumption of a monospecific composition of the Polzberg coleoid fauna. Instead, we report the occurrence of the new phragmoteuthid Phragmoteuthis polzbergensis nov. sp. and a newly excavated specimen of Phragmoteuthis indicates the presence of ten arms within the group of the Phragmoteuthida for the first time. Phragmocones with small opening angles combined with cylindrical (roundly closed) body chambers, and arm hook types which are unusual for phragmoteuthids indicate the presence of the basal coleoid group comprising the rostrum-less genus Mojsisovicsteuthis.
... As described in previous papers 4,20,25 , the lowermost, fossiliferous part of the Reingraben formation can be dated based on the occurrence of Austrotrachyceras minor, Paratrachyceras haberfellneri and the accompanying Carnites floridus and Simonyceras simonyi, indicating a Julian 2 age (Julian 2/Ib; Fig. 2). The presence of these ammonoid members together with the mass occurrence of the benthic bivalve Halobia rugosa within the fossiliferous layers confirms the biostratigraphical age assignment as Julian 4,8,20,21,25 . ...
... The concentration is low in Göst Top, high near the base of the Reingraben formation and decreases upwards within the latter unit. The ratio between C 13-17 and C [18][19][20][21][22] aryl isoprenoids (aryl isoprenoid ratios (AIR 52 ) is typically low (< 0.21). In contrast to aryl isoprenoids, isorenieratane was not detected. ...
... Although this may reflect temporally different redox conditions in different portions of the water column, these results should not be over-interpreted. Nonetheless, the laminated sediments were clearly deposited under oxygen-depleted conditions 20,21,24 . The oxygen depletion was probably related to salinity stratification 20,21,24 . ...
We present a multi-proxy investigation of a lower Carnian basinal succession from Polzberg in the Northern Calcareous Alps (Lower Austria). A section comprising a unique Konservat-Lagerstätte was studied based on bio- and chemostratigraphy along with geophysical methods, yielding a detailed and robust stratigraphic calibration of the Polzberg succession. The Polzberg section revealed the paleoceanographic history and helped to identify a global climatic reversal, the Carnian Pluvial Episode. The age of the Upper Triassic Reingraben formation in the Northern Calcareous Alps is refined as the Austrotrachyceras austriacum Zone within the lower Carnian (Julian 2). Ammonoids and conodonts provide a detailed biostratigraphic subdivision that serves as a basis for analyses of the faunal distribution and the paleoenvironmental evolution of the Upper Triassic Reifling Basin. The succession includes lithological and facies changes similar to those of coeval units in the Tethys. The Carnian was characterized by a weak (~ 1‰) positive δ¹³C trend, punctuated by a negative shift during the lower Carnian corresponding to the initiation of the Carnian Pluvial Episode, a period representing the onset of early/late Carnian transitional global greenhouse conditions. Organic maturity parameters and the conodont alteration index (CAI) show that the thermal overprint of the Polzberg section is low. Biomarker proxies suggest that the organic matter of the uppermost Göstling formation is a mixture of marine and terrestrial material deposited in a dysoxic environment. Within the overlaying Reingraben formation, the amount of marine biomass decreased gradually upwards. Oxygen-depleted conditions, probably due to water-column stratification, continued during deposition of the Reingraben formation. Bacterial sulfate reduction played a major role in organic matter degradation.
... Aulacoceratid rostra (Aulacoceratida Stolley, 1919) are more abundant, but mostly without further information on their phragmocone and soft tissue morphology. Records of gladius-bearing coleoids are very rare and not beyond any doubts (Fuchs, 2020;Lukeneder & Lukeneder, 2022;Reitner, 1978). Belemnitid belemnoids probably arose during the Triassic, but so far only a handful of rostra are described (Iba et al., 2012). ...
... The order Phragmoteuthida Mojsisovics, 1882, a Triassic-Jurassic group that links aulacoceratids with a long tubular body chamber and Mesozoic belemnitids with a slender spatulate proostracum are comparatively well known (Doguzhaeva et al., 2007;Fuchs & Donovan, 2018;Lukeneder & Lukeneder, 2022). Earliest records of phragmoteuthids are known with question marks from the Upper Permian, but phragmocones with their characteristic broad three-lobed proostracum certainly appear in the Middle Triassic (Anisian). ...
Two stratigraphically well constrained (by ammonites and conodonts) coleoid remains have been recorded from the Triassic (Anisian) dark-grey organodetritic limestones (Ráztoka Limestone) of Western Carpathians (Hronic Nappe). The limestones deposited at the periphery of a former carbonate platform. It yields a highly diverse cephalopod fauna including nautiloids (2 taxa), ammonoids (7 taxa) and indetermined aulacoceratids. Two unusual coleoid specimens are referred to genus Mojsisovicsteuthis ( M . boeckhi ) and probably to a new taxon (described as Breviconoteuthis aff. breviconus herein) possessing similar morphological features of genus Breviconoteuthis (Phragmoteuthida) and/or Zugmontites . Based on index ammonites and conodonts, both records are of the uppermost Trinodosus through the lowermost Reitzi zones (Anisian—lower Illyrian). While the genus Mojsisovicsteuthis has been widely dispersed (however its records are rare), the occurrence of Breviconoteuthis and Zugmontites is strictly limited to the Alpine-Carpathian region. Comparing with the holotype and additional specimens stored in the Hungarian Natural History Museum, the overal shell of Mojsisovicsteuthis and its size has been reconstructed. Its relationship to aulacoceratids and phragmoteuthids is briefly discussed. Geochemical record ( n -alkanes from the bulk rock) provided a relevant signal of the existence of algal meadows.
... Aulacoceratid rostra (Aulacoceratida Stolley, 1919) are more abundant, but mostly without further information on their phragmocone and soft tissue morphology. Records of gladius-bearing coleoids are very rare and not beyond any doubts (Reitner, 1978, Fuchs 2020, Lukeneder & Lukeneder 2022). Belemnitid belemnoids probably arose during the Triassic, but so far only a handful of rostra are described (Iba et al. 2012). ...
... The order Phragmoteuthida Mojsisovics, 1882, a Triassic -Jurassic group that links aulacoceratids with a long tubular body chamber and Mesozoic belemnitids with a slender spatulate proostracum are comparatively well known (Doguzhaeva et al. 2007, Fuchs et Donovan, 2018Lukeneder & Lukeneder 2022). Earliest records of phragmoteuthids are known with question marks from the Late Permian, but phragmocones with their characteristic broad three-lobed proostracum certainly appear in the Middle Triassic (Anisian). ...
The two stratigraphically well constrained (by ammonites and conodonts) coleoid remains have been recorded in the Triassic (Anisian) dark-grey organodetritic limestones (Ráztoky Limestone) of Western Carpathians (Hronic Nappe). The facies originated between carbonate ramp and basinal development. It yields highly diversificated cephalopod fauna including nautiloids (2 taxa), ammonoids (7 taxa) and indetermined aulacoceratids. Two unusual coleoid specimens are referred to genus Mojsisovicsteuthis ( M . boeckhi ) and probably to a new taxon (described as Breviconoteuthis aff. breviconus herein) possessing similar morphological features of genus Breviconoteuthis (Phragmoteuthida) and/or Zugmontites . Based on index ammonites and conodonts, both records are of the uppermost Trinodosus through the lowermost Reitzi zones (Anisian – lower Illyrian). While the genus Mojsisovicsteuthis has been widely dispersed (however its records are rare), the occurrence of Breviconoteuthis and Zugmontites is strictly limited to the Alpine-Carpathian region. Compare to the holotype and additional specimens stored in the Hungarian Natural History Museum, the overal shell of Mojsisovicsteuthis and its size has been reconstructed. Its relationship to aulacoceratids and phragmoteuthids is briefly discussed. Geochemical record provided a relevant signal of the existence of algal meadows which may represent an analogy with later sea-grasses.
... A. Lukeneder , P. Lukeneder mineralized belemnoid cartilage (Lukeneder & Lukeneder, 2022b). These publications were based on historical and recent palaeontological material housed in the official Austrian collections and material from private collections. ...
... A. minor and P. haberfellneri appear frequently with partially preserved buccal apparatuses of anaptychus-type lower jaws (Trauth, 1935) and upper jaws (Lukeneder & Lukeneder, 2022a). In numerous specimens of Phragmoteuthis bisinuata (Doguzhaeva & Summesberger, 2012;Doguzhaeva et al., 2007b;Lukeneder & Lukeneder, 2022b), the tripartite proostracum and the phragmocones appear with black bituminous sheets of the ink sac, along with black amorphous cartilage and arm hook structures (Lukeneder & Lukeneder, 2021, 2022b. For the questionable belemnoid Lunzoteuthis schindelbergensis from the same locality see Doguzhaeva et al. (2006). ...
... A. minor and P. haberfellneri appear frequently with partially preserved buccal apparatuses of anaptychus-type lower jaws (Trauth, 1935) and upper jaws (Lukeneder & Lukeneder, 2022a). In numerous specimens of Phragmoteuthis bisinuata (Doguzhaeva & Summesberger, 2012;Doguzhaeva et al., 2007b;Lukeneder & Lukeneder, 2022b), the tripartite proostracum and the phragmocones appear with black bituminous sheets of the ink sac, along with black amorphous cartilage and arm hook structures (Lukeneder & Lukeneder, 2021, 2022b. For the questionable belemnoid Lunzoteuthis schindelbergensis from the same locality see Doguzhaeva et al. (2006). ...
Here we report new data on the Upper Triassic Polzberg Konservat-Lagerstätte in the Northern Calcarous Alps of Lower Austria. We examined new fossil material obtained from bed-by-bed collections of the well-laminated Reingraben Shales. Over 5290 new fossils of various marine taxa were collected within the fossiliferous layers from the Early Carnian (Julian 2 Ib, Austrotrachyceras austriacum Zone). The newly collected assemblage comprises ammonoids ( Austrotrachyceras , Paratrachyceras , Carnites , Simonyceras ), belemnoids ( Phragmoteuthis , Lunzoteuthis ), bivalves ( Halobia , div. indet taxa), gastropods (caenogastropods/heterobranchs), thylacocephalan arthropods ( Austriocaris, Atropicaris ), crustaceans (the decapod Platychela and isopods Obtusotelson , Discosalaputium ), branchiopods ( Euestheria ), polychaetes ( Palaeoaphrodite sp. and an unidentified eunicid polychaete), acytinopterygians ( Saurichthys , Polzbergia, Peltopleurus, Habroichthys ), cartilaginous fishes ( Acrodus ), coelacanth fish ( “Coelacanthus” ), a lungfish ( Tellerodus ), and numerous conodont clusters ( Mosherella ). Bromalites (coprolites and regurgitalites) produced by piscivorous actinopterygians and durophagous fish accompany the Polzberg palaeobiota along with rare plant remains ( Voltzia, div. indet plants). Based on new findings the palaeobiota characterises an intermittent colonisation by abundant benthic halobiid bivalves and a predator-dominated (fish, belemnoids) nektic community. The prerequisites for high-quality preservation—calm and oxygen-depleted conditions—prevailed at the sea floor of the Polzberg Konservat-Lagerstätte . Normal marine conditions prevail in the Reifling Basin, occasionally interrupted by freshwater influx. New in situ findings of benthic and nektic taxa highlight the great value of the unique Polzberg palaeobiota and the autochthonous deposition of the inhabitants within the palaeohabitat. The fauna and flora from the Polzberg Konservat-Lagerstätte , deposited during the Carnian Pluvial Episode or Carnian Wet Intermezzo, points to a carbonate platform decline followed by the deposition of laminated deposits in warmer and wetter conditions.
... Lukeneder and Lukeneder (2021) preliminarily determined the black structures from Polzberg as "teuthid cartilages". A detailed comparison of 64 of these seemingly amorphous structures from Polzberg (n = 57) and Cave del Predil (n = 7) has meanwhile shown that the structures from Polzberg and Cave del Predil are identical, because they respectively consist of a pair of open rings with wing-like extensions (Lukeneder and Lukeneder 2022). ...
... On the histological level, there are strong similarities between the cartilage of Recent cephalopods and vertebrates (Bairati et al. 1987;Cole and Hall 2004). We therefore scrutinized the ultrastructure of 13 of these black, multi-elemental structures using various techniques (micro-CT, SEM, EDS, thin sections; Lukeneder and Lukeneder 2022). These ultrastructural analyses combined with taphonomic investigations on numerous fossil taxa of the Polzberg palaeobiota as well as actuo-palaeontological comparisons with extant cephalopods (species of Sepia, Loligo, Octopus) revealed a well preserved, branching canal system that is typical for coleoid cartilage (Lukeneder and Lukeneder 2022 and references therein; PL, AL, Christina Kaurin, and Valentin Blümel, unpublished data). ...
... We therefore scrutinized the ultrastructure of 13 of these black, multi-elemental structures using various techniques (micro-CT, SEM, EDS, thin sections; Lukeneder and Lukeneder 2022). These ultrastructural analyses combined with taphonomic investigations on numerous fossil taxa of the Polzberg palaeobiota as well as actuo-palaeontological comparisons with extant cephalopods (species of Sepia, Loligo, Octopus) revealed a well preserved, branching canal system that is typical for coleoid cartilage (Lukeneder and Lukeneder 2022 and references therein; PL, AL, Christina Kaurin, and Valentin Blümel, unpublished data). Doguzhaeva et al. (2022: figs. ...
Marine conservation deposits (‘Konservat-Lagerstätten’) are characterized by their mode of fossil preservation, faunal composition and sedimentary facies. Here, we review these characteristics with respect to the famous conservation deposit of the Besano Formation (formerly Grenzbitumenzone; including the Anisian–Ladinian boundary), and the successively younger fossil-bearing units Cava inferiore, Cava superiore, Cassina beds and the Kalkschieferzone of Monte San Giorgio (Switzerland and Italy). We compare these units to a selection of important black shale-type Lagerstätten of the global Phanerozoic plus the Ediacaran in order to detect commonalities in their facies, genesis, and fossil content using principal component and hierarchical cluster analyses. Further, we put the Monte San Giorgio type Fossillagerstätten into the context of other comparable Triassic deposits worldwide based on their fossil content. The results of the principal component and cluster analyses allow a subdivision of the 45 analysed Lagerstätten into four groups, for which we suggest the use of the corresponding pioneering localities: Burgess type for the early Palaeozoic black shales, Monte San Giorgio type for the Triassic black shales, Holzmaden type for the pyrite-rich black shales and Solnhofen type for platy limestones.
Although patchy, the fossil record of coleoids bears a wealth of information on their soft part anatomy. Here, we describe remains of the axial nerve cord from both decabrachian (Acanthoteuthis, Belemnotheutis, Chondroteuthis) and octobrachian (Plesioteuthis, Proteroctopus, Vampyronassa) coleoids from the Jurassic. We discuss some hypotheses reflecting on possible evolutionary drivers behind the neuroanatomical differentiation of the coleoid arm crown. We also propose some hypotheses on potential links between habitat depth, mode of life and the evolution of the Coleoidea.
Supplementary Information
The online version contains supplementary material available at 10.1186/s13358-023-00285-3.
Wetlands are important to continental evolution, providing both arenas and refugia for emerging and declining biotas. This significance and the high preservation potential make the resulting fossiliferous deposits essential for our understanding of past and future biodiversity. We reconstruct the trophic structure and age of the early Permian Manebach Lake ecosystem, Germany, a thriving wetland at a time when the tropical biosphere faced profound upheaval in the peaking Late Palaeozoic Icehouse. Nine excavations, high-resolution spatiotemporal documentation of fossils and strata, and U-Pb radioisotopic dating of tuffs allow us to distinguish autogenic and allogenic factors shaping the limnic biocoenosis. The Manebach Lake was an exorheic, oxygen-stratified, perennial water body on the 10¹–10² km² scale, integrated into the catchment draining much of the European Variscides. Lake formation paralleled an Asselian regional wet climatic interval and benefited from rising base level due to post-Variscan half-graben tectonics. Stromatolite-forming cyanobacteria, bivalves, several crustaceans , amblypterids and xenacanthid sharks formed a differentiated biocoenosis in the lake. Fossil stomach remains and teeth prove the rare presence of acanthodians, branchiosaurs and large amphibians. The results indicate woody-debris-bearing lake littorals devoid of semi-aquatic and aquatic plants as places suitable for stromatolites to grow, underpin the model of declining freshwater-shark diversity in most Permian Variscan basins, demonstrate fish/amphibian ratios in limnic assemblages to measure lake perenniality and reveal taphonomic biases in lake taphocoenoses. Our outcomes call for more knowledge about the diversity, ecology and fossilization pathways of past limnic biotas, particularly microorganisms and actinopterygian fishes, to reconstruct deep-time continental ecosystems.
