Metriorhynchids are the only crocodyliforms adapted to pelagic marine life. Snout natural endocasts of the Tithonian (Late Jurassic) metriorhynchid Cricosaurus araucanensis indicated that skeletal changes defining the peculiar metriorhynchid body plan were coupled with changes of the soft cephalic anatomy such as the enlarged salt glands and restructuring of the paranasal sinus system. Seven new natural endocasts of the snout and a 3-D reconstruction of C. araucanensis are described. Data from these casts and the reconstruction are congruent, and they are combined into an accurate reconstruction that improves our knowledge of the pre-orbital anatomy. The olfactory tract, bulbs, olfactory nasal region and the anterior extension of the antorbital sinus within the maxilla are recognized. Osteological correlates of the salt gland body are also proposed. Palaeobiological inferences are erected based on the integration of natural endocasts and 3-D reconstruction data. It is proposed that C. araucanensis nasal salt glands were highly vascularized with a blood supply comparable with those of extant marine birds. Reduced olfactory bulbs and olfactory nasal region indicate that the aerial olfaction, differing from extant crocodilians, was not well developed.
]]>In this work, we analyse cheek teeth crown complexity through the calculation of fractal dimension in the giant caviomorph rodent Eumegamys paranensis (Late Miocene of Argentina) and evaluate its functional significance. Our results indicate that, in all teeth of Eumegamys paranensis, the fractal dimension was around 1.5, similar to the Koch quadratic curve type two. The anterior portions of the molars, with the highest values of fractal dimension, are interpreted as areas that supported greater occlusal pressures. Crown complexity in E. paranensis is related to the increased mechanical capacity to process relatively demanding food items and to allow more food to be divided in each masticatory cycle. Eumegamys paranensis would have been a mixed feeder, consuming a variable diet obtained close to the ground. This feeding behaviour is compatible with the heterogeneous environment inferred for the Mesopotamic area during the Late Miocene. E. paranensis was probably a wide ranging species, being able to eat close to water bodies and in gallery forests that occurred in the surrounding of the pre-Paraná river system.
]]>The shell repair scars of modern Nautilus pompilius in the Philippines presumably due to sub-lethal predatory attacks were examined throughout ontogeny ranging from hatching to maturity, revealing the higher risk of fatality in earlier ontogenetic stages. The examinations throughout ontogeny demonstrate that: (1) sub-lethal predatory attacks by other organisms have no preferred position through ontogeny; (2) the sizes of shell repair scars are similar throughout ontogeny and, therefore, the width and length of shell repair scars relative to shell diameter decrease from hatching towards maturity; and (3) the proportion of sub-lethal predatory attacks within 10 mm increments of shell diameter are similar irrespective of shell diameter or decrease approaching maturity. The numbers of the irregular, radiating, black shell repair scars, indicating injury to the soft parts at shell breakage, decrease at a diameter >80 mm and none were found at <56 mm in shell diameter. Furthermore, the rarity of juvenile trapping data might be related to the higher fatality from predatory attacks in earlier ontogenetic stages, providing crucial information for the conservation of N. pompilius.
]]>The Frasnian–Famennian extinction witnessed the global devastation of both level-bottom and reef communities in low latitudes. Marine extinctions in offshore level-bottom communities are associated with two widespread, transgressive, anoxic ‘Kellwasser Events’ that support an anoxia–extinction link. Typical Kellwasser facies of bituminous limestones and shales are not obviously recorded in shallow-water settings, and thus, it is unclear whether anoxia played a role in reef losses. We evaluate geochemical, petrographic and facies evidence for oxygen restriction from an extremely shallow-water carbonate platform in Alberta. Sequence stratigraphy places the Frasnian–Famennian boundary at a sequence boundary that tops a laminated mudstone and interrupts carbonate platform deposition. Two transgressive pulses have been identified, one of which is associated with the second, major transgression of T-R cycle IId of the Devonian eustatic sea-level curve. Geochemical proxies indicate that these transgressions were accompanied by influx of dysoxic or anoxic waters. Organic carbon and U enrichment in the Frasnian, particularly just below the Frasnian–Famennian boundary, points to episodic dysoxic conditions that probably persisted into the basal Famennian and were coincidental with the global Upper Kellwasser Event. This study provides the first evidence for the smoking gun of an anoxia-driven extinction in very shallow waters, implicating this potent killer in the demise of the Devonian reefs.
]]>The distinctive brachiopod Dicoelosia King 1850 is characterized by a strongly bilobed outline. To date, studies have concentrated on its functional morphology, taxonomy and evolution; little attention has been paid to its ontogeny. Here, we map population variation by principal component analysis for over 80 specimens distributed across five species of Dicoelosia. Using geometric morphometrics with landmarks for some 40 specimens, the ontogenic trends in D. sp. nov. are compared with those of Dicoelosia biloba. In addition, the ontogenic pathway in D. sp. nov. is investigated by morphing with control points, a new technique introduced here to palaeontology. Combining the results above, the ontogeny of the key character of the genus, emargination, is modelled. Within single populations, taxa may develop from broad weakly emarginate forms into those that are elongate and deeply emarginate. As the identification of the genus and its species depends on external morphological characters, the definition of ontogenetic trends in each species is essential for taxonomic discrimination. Substantial population variation exists in many of its species; however, the morphing technique provides a method of simulation, predicting the full range of ontogenetic variation in given populations.
]]>The Middle Eocene Man Aike Formation of southern Patagonia, Argentina, offers the opportunity to study the palaeoecological and palaeoenvironmental significance of Macaronichnus ichnofabrics in a tide-dominated, coastal embayment compound-dune complex. Four recurrent Macaronichnus ichnofabrics are recognized: Macaronichnus IF-1 characterized by Macaronichnus segregatis segregatis; Macaronichnus IF-2 with M. segregatis segregatis, Ophiomorpha nodosa, Skolithos linearis and rare Palaeophycus tubularis; Macaronichnus IF-3 typified by M. segregatis segregatis, O. nodosa, Asterosoma radiciforme and rare S. linearis; and Macaronichnus IF-4 consisting of M. segregatis segregatis, Nereites missouriensis, A. radiciforme, O. nodosa, Rosselia socialis, S. linearis, P. tubularis, Planolites isp. and Chondrites isp. Variations in these ichnofabrics are linked to changes in local hydraulic conditions in the upper and lower trough areas of the compound dunes. In high-energy, shallow and marginal marine environments, the Macaronichnus tracemaker is commonly the first to colonize the shifting, sandy substrates. Overprinting by other ichnoguilds occurs only after the physiochemical conditions became favourable and the colonization window length increases.
]]>Chan, N.R., Dyke, G.J. & Benton, M.J. 2013: Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds. Lethaia, Vol. 46, pp. 146–152.
Although many Mesozoic fossil birds have been found with primary feathers preserved, these structures have rarely been included in morphometric analyses. This is surprising because the flight feathers of modern birds can contribute approximately 50% of the total wing length, and so it would be assumed that their inclusion or exclusion would modify functional interpretations. Here we show, contrary to earlier work, that this may not be the case. Using forelimb measurements and primary feather lengths from Mesozoic birds, we constructed morphospaces for different clades, which we then compared with morphospaces constructed for extant taxa classified according to flight mode. Consistent with older work, our results indicate that among extant birds some functional flight groups can be distinguished on the basis of their body sizes and that variation in the relative proportions of the wing elements is conservative. Mesozoic birds, on the other hand, show variable proportions of wing bones, with primary feather length contribution to the wing reduced in the earlier diverging groups. We show that the diverse Mesozoic avian clade Enantiornithes overlaps substantially with extant taxa in both size and limb element proportions, confirming previous morphometric results based on skeletal elements alone. However, these measurements cannot be used to distinguish flight modes in extant birds, and so cannot be used to infer flight mode in fossil forms. Our analyses suggest that more data from fossil birds, combined with accurate functional determination of the flight styles of living forms is required if we are to be able to predict the flight modes of extinct birds. □Birds, flight, morphospace, Mesozoic, wing.
Krapovickas, V., Mancuso, A.C., Marsicano, C.A., Domnanovich, N.S. & Schultz, C.L. 2013: Large tetrapod burrows from the Middle Triassic of Argentina: a behavioural adaptation to seasonal semi-arid climate? Lethaia, Vol. 46, pp. 154–169.
We report the discovery of large burrow casts in the early Middle Triassic Tarjados Formation, at Talampaya National Park, north-western Argentina. Facies analysis indicates the burrows are preserved in sandbars deposited by an ephemeral river under semi-arid and seasonal climatic conditions. The structures are mostly preserved in longitudinal cross-section and consist of an opening, an inclined tunnel (ramp), and a terminal chamber. The ramp is 8–14 cm in height, up to 130 cm in length and penetrates 49–63 cm bellow the palaeosurface with an inclination of 22°–30°. We studied burrow cast dimensions, overall architectural morphology, surficial marks, and compared them with other large burrows of both invertebrate and vertebrate origin. A tetrapod origin of the burrow casts was established based on: distinctive architecture, and size, which is more than twice the most common size range for large terrestrial invertebrate burrows. Comparison with other Upper Permian and Triassic tetrapod burrows allows us to identify three general morphological groups: (1) simple inclined burrows; (2) helical burrows; and (3) burrow network complexes, representing different behaviours. A study of tetrapod body fossils preserved within other Upper Permian and Triassic burrows shows that the Tarjados structures were most likely produced by non-mammalian cynodonts. The environmental and climatic context suggests that aridity and seasonality played a fundamental role selecting burrowing behaviour in therapsids and that by the Early–Middle Triassic their burrowing behaviour attained a complexity comparable to modern mammals. □Argentina, behaviour, palaeoclimate, Permo-Triassic, Tarjados Formation, Tetrapod burrows.
Benedetto, J.L. 2013: Presence of punctae in the ‘plectorthoidean’ brachiopod Famatinorthis turneri (Middle Ordovician) from western Argentina: implications for early diversification of punctate orthides. Lethaia, Vol. 46, pp. 170–179.
Famatinorthis Levy & Nullo is a distinctive orthide brachiopod of Dapingian age from the volcaniclastic rocks of the Famatina Range of western Argentina. Although it was originally classified among the plectorthoideans, a new collection from the La Escondida Formation has yielded exceptionally well-preserved moulds of Famatinorthis turneri in which silicified infillings of punctae are clearly visible, leading to the reassignment of the genus to the dalmanellidines. In this paper, phylogenetic analyses are used to determine the evolutionary relationships of Famatinorthis, the Tremadocian linoporellid Lipanorthis, and other Ordovician Gondwanan genera. The placement of Plectorthoidea in the same major clade as linoporellids, and the separation of Dalmanellidina as an independent clade are the most important features of all shortest trees, supporting the idea that linoporellids may have originated from a plectorthoid ancestor. Cladistic analysis reveals that Lipanorthis lies close to the ancestry of the linoporellid lineage, and Famatinorthis clusters within the more derived taxa of the clade with which it shares a large septalium. It seems that the presence of endopunctae in the orthides does not necessarily indicate close phylogenetic relationships as it could have occurred at different times in different clades. If the homoplasic nature of endopunctae in the order Orthida is supported by further morphologic and phylogenetic studies, the fundamental division of orthides in non-punctate (Orthidina) and punctate (Dalmanellidina) may need revision. □Brachiopods, Ordovician, Gondwana, Famatina, phylogeny, punctate orthides.
Mosasaurs were common predators on the ammonites that inhabited the upper water column of the Late Cretaceous Western Interior epicontinental seaway of North America. Mosasaurs developed predictable behaviour patterns for feeding on ammonite prey. There are no previous reports of mosasaur predation on the much less common Cretaceous nautiloids, possibly because of the prey's predominantly deep, epibenthic habitat, as deduced from modern Nautilus life habits. A single specimen of the highly inflated nautiloid, Eutrephoceras dekayi (Conrad), prey to a small adult mosasaur, likely Platycarpus, Prognathodon or Mosasaurus, is reported herein from the Pierre Shale of Colorado in the Early Maastrichtian biozone of Baculitesgrandis transitional to the biozone of B. clinolobatus. The nautiloid was killed in the same manner as described previously for discoid ammonites (Placenticeras, Sphenodiscus) from coeval strata in the USA and Canada.
]]>Multivariate analysis based on nine biometric characters of 171 Late Ordovician rhynchonellide specimens from nine upper Sandbian–upper Katian localities in North America supports the hypothesis that one of the diagnostic taxa of the North American epicontinental brachiopod fauna, Hiscobeccus, evolved from Rhynchotrema, which lived predominantly in peri-cratonic settings. The oldest known Hiscobeccus, H. mackenziensis of early Katian age, exhibits transitional characteristics between Rhynchotrema and Hiscobeccus, and it clusters more closely with Rhynchotrema than with younger species of Hiscobeccus of mid–late Katian (Maysvillian–Richmondian) age. Diversification of the Hiscobeccus lineage in epicontinental seas was characterized by drastic increase in shell size, globosity and lamellosity, especially in palaeoequatorially located inland basins. Such morphological trends are interpreted as adaptation to relatively shallow, muddy substrates, moderate water turbulence, relatively low oxygen content and unstable supply of nutrients in generally overheated epicontinental seas with sluggish circulation.
]]>Fossilized ring-like structures with enigmatic function and taxonomic affiliation were recovered for the first time from the Upper Ordovician of the Carnic Alps and the Silurian of Bohemia. These rings, already mentioned as minor constituents in previous conodont studies (e.g. Webers 1966, p. 1; Bischoff 1973, p. 147), were reported from the Palaeozoic of several regions in Europe and North America. Originally considered as inwardly accreted adhering discs of a benthic hyolithelminth worm with a phosphatic tubular projection, they were later reinterpreted in relation to a putative crinoid epibiont or even as possible scyphozoans. Despite a long debate, neither the function of the enigmatic Palaeozoic rings nor their taxonomic affiliation has been fully clarified. The studied material, extracted by a standard technique in use for conodonts, consists of 235 elements from 16 stratigraphic levels in the Plöcken Formation (Carnic Alps, Cellon Section; Amorphognathus ordovicicus Biozone, Hirnantian, Ordovician) and in the Kopanina Formation (Bohemia, Mušlovka Quarry; Polygnathoides siluricus Biozone, Ludfordian, Silurian). To explore whether ring size and shape changed over time, we employed a novel combination of geometric morphometric approaches for outlines with no ‘homologous’ landmarks and showed that only size appreciably varied with an increase of ca. 20%. The emerging data from this study are consistent with the interpretation of the rings as an adhering structure of a benthic organism living on a relatively uniform hard substrate.
]]>X-ray microtomography (XMT) is a non-invasive and non-destructive method that has often been used to study fossils. It allows serial sections to be made as little as few micrometers apart; such a resolution is unachievable for classical serial sectioning; moreover, in contrast to the latter, the specimen is not destroyed. Microtomography can, however, be applied only in cases where differences in X-ray absorption between the skeleton and its infilling are great. We show that this method may be also applied to tabulate corals. Case studies of blastogeny are based on Silurian (Aulopora, Favosites) and Devonian (Thamnopora) specimens from Poland. We show that the sequence of events in the blastogeny of Aulopora sp. is different from that of ‘Aulopora serpens minor’ from the Devonian of the Holy Cross Mountains and similar to auloporids from the Devonian of England. Blastogeny in Favosites is very similar to that known from the related genera Squameofavosites and Thamnopora. This suggests that members of the genus Aulopora may be more diversified within the genus (as presently understood) than genera within the Favositidae.
]]>Marine sessile benthic organisms living on hard substrates have evolved a variety of attachment strategies. Rhizotrochus (Scleractinia, Flabellidae) is a representative azooxanthellate solitary scleractinian coral with a wide geographical distribution and unique attachment structures; it firmly attaches to hard substrates using numerous tube-like rootlets, which are extended from a corallum wall, whereas most sessile corals are attached by stereome-reinforced structures at their corallite bases. Detailed morphological and constructional traits of the rootlets themselves, along with their evolutionary significance, have not yet been fully resolved. Growth and developmental processes of spines in Truncatoflabellum and rootlets in Rhizotrochus suggest that these structures are homologous, as they both develop from the growth edges of walls and are formed by transformation of wall structures and their skeletal microstructures possess similar characteristics, such as patterns of rapid accretion and thickening deposits. Taking molecular phylogeny and fossil records of flabellids into consideration, Rhizotrochus evolved from a common free-living ancestor and invaded hard-substrate habitats by exploiting rootlets of spines origin, which were adaptive for soft-substrate environments.
]]>The bioeroding foraminifer Troglotella incrustans Wernli and Fookes (Bolletino della Societa Paleontologica Italiana 31, 1992, 95), is widely reported from Bajocian?, and Oxfordian to Lower Cenomanian (with a Late Jurassic acme) shallow-water limestones of the Tethyan realm. A single specimen of a boring foraminifer, assigned to T. incrustans, has now been observed from the Lower Permian (Sakmarian) Community Pit Formation of the Doña Ana Mountains, New Mexico, USA. Surviving the end-Permian mass extinction, T. incrustans might be a Lazarus taxon that persisted in refuges. This finding represents the oldest record of a foraminifer exhibiting an euendolithic way of life. Boring foraminifera have not been previously recorded from strata older than the Jurassic. Boring traces of potentially foraminiferan origin, however, have been already reported from the Lower Carboniferous (?Ordovician).
]]>A detailed ichnological analysis of the Upper Devonian–Lower Mississippian Bakken Formation of sub-surface Saskatchewan and the partially coeval Exshaw Formation of Alberta indicates the presence of an anomalous ichnofacies gradient. The distal Cruziana Ichnofacies, which in rocks of other ages is restricted to lower-offshore facies, here ranges from this setting to the lower shoreface. No archetypal Cruziana Ichnofacies is present in these deposits. This pattern is interpreted as resulting from the differential effects of the Late Devonian mass extinction in shallow-water ecosystems. The onshore expansion evidenced by ichnological data is consistent with the pattern displayed by the body-fossil record, which indicates a re-invasion of shallow-water environments by the Palaeozoic evolutionary fauna during the Late Devonian and into the Early Carboniferous. The ichnofauna studied is overwhelmingly dominated by deposit feeders, with suspension feeders being notably absent, further underscoring the importance of trophic type as a selectivity trait during mass extinctions.
]]>A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.
]]>An isolated track from the latest Campanian–Maastrichtian Wapiti Formation (Alberta) is tentatively identified as a pterosaur manus print. The basic digital formula (digit I < digit II < digit III) is consistent with the plesiomorphic condition in pterosaurs. The track measures 25.5 cm in length, making it the largest putative pterosaur manus print (estimated wingspan 7.7 m) from North America. Although precise chronostratigraphic data are lacking, sedimentary evidence indicates the track comes from a mid- to late Campanian fluvial deposits that accumulated approximately 400 km from the closest shoreline within a high-latitude (65°N) setting.
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