Lakes and rivers of the younger Paleozoic and as well in the Triassic could not only house tetrapod and bony fish vertebrate dwellers but also selachian predators, in particular the Xenacanthida, well known for their characteristic neck spines, and the Hybodontiformes, which display a pair of lateral head spines and characteristic fin spines. The latter are distinct from those of the Acanthodii (popularly also referred to as "spiny sharks"), a group of basal vertebrates that ocurred in freshwater environments as well, but became extinct before the beginning of the Triassic.
Complete shark specimens are seldom recorded, the same is true for complex finds comprising a couple of skeletal elements from the same individual - taxonomists often have to deal with assemblages of individual scales, spines, and teeth and systematics heavily relies on tooth characteristics (e.g. Schneider 1988 for the Xenacanthida, Rees 2008 for hybodont sharks).
Like the recent bullhead sharks (Heterodontus) at least some of the Carboniferous to Triassic freshwater sharks were oviparous - different types of spiral egg capsules not quite unlike those capsules of Heterodontus occur in different types of freshwater environments, e.g. marginal lake sediments or low-energy river banks; often they appear unrelated to skeletal remains. This has been interpreted as being indicative for a separation between the actual habitats of the sharks and their spawning grounds (about the facial aspects: see Schneider & Reichel 1989). To what extent the occurrence of xenacanth and hybodont sharks in freshwater deposits is indicative for a marine influence is currently a matter of debate.
In the first descriptions of the 19th century fossil egg capsules were misinterpreted as cone-like fructifications of some kind of plant. This was due to the rhomboidal pattern the egg capsule impressions often display as consequence of taphonomic flattening (and the consequent overlap of the spiral patterns on the front and back sides). Two types of shark egg capsules have been recovered from the Madygen Formation during fieldwork in 2007 (Fischer et al. 2007) - more on that later.
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Jan Fischer, featured in the last FPhotW, who is working on the Madygen chrondrichthyans (as soon as they appear) and isotope paleontology of shark teeth/spines, is thanked here for supplying me with literature. (Hopefully I can convince Jan to write a guest contribution).
Refs:
Synoptical papers:
Maisey, J.G. (1982): The Anatomy and Interrelationships of Mesozoic Hybodont Sharks. - American Museum Novitates 2724: 1- 48; New York.
Schneider, J. W. & Zajic, J. (1994): [Xenacanths (Pisces, Chondrichthyes) of the middle European Upper Carboniferous and Permian - revision of the originals of GOLDFUSS 1847, BEYRICH 1848, KNER 1867 and FRITSCH 1879-1890.] - Freiberger Forschungshefte, C 452: 101-151; Leipzig.
On tooth systematics:
Schneider, J.W. (1988): [Basics of the morphogeny, taxonomy, and biostratigraphy of isolated xenacanth teeth (Elasmobranchii)]. - Freiberger Forschungshefte, C 419: 71- 80; Leipzig.
Rees (2008): Interrelationships of Mesozoic hybodont sharks as
indicated by dental morphology – preliminary results. - Acta Geologica Polonica 58 (2): 217-221.
On egg capsules:
Schneider, J.W. & Reichel, W. (1989): [Chondrichthyan egg capsules from the Rotliegend (Lower Permian) of Middle Europe - conclusions regarding the palaeobiogeography of palaeozoic freshwater sharks.] - Freiberger Forschungshefte, C 436: 58- 69; Leipzig.
Fischer, J., Voigt, S. & Buchwitz, M. (2007):
First elasmobranch egg capsules from freshwater lake deposits of the Madygen Formation (Middle to Late Triassic, Kyrgyzstan, Central Asia). - Paläontologie, Stratigraphie, Fazies (15), Freiberger Forschungshefte, C 524: 41-46; Freiberg.
Freitag, 23. Januar 2009
Mittwoch, 21. Januar 2009
Fieldwork Photo of The Week
Morning time at the digging site: Having contemplated the vast landscape bare of human presence but sprenkled with the litter of a forgotten civilization graduate student Jan Fischer decides on the right moment for starting his daily business of not moving to many a rock at once and feigning the impression of having survived another day in this vale of tears only at the close of livelong fatigue and exhaustion...
Mittwoch, 7. Januar 2009
Fieldwork Photo of the Week
See the spy in the back? Some Kyrgyz aborigines can't help being distrustful...I'm thankful to Juliane Hentschke who safed us these moments and a number of bones.
Samstag, 3. Januar 2009
Tectonics & Paleo (4):
The world of CPOs and ODFs
Materials in geoscience and biology are often not isotropic - their properties, such as conductivity, soundwave velocity, and shear strength, vary with direction.
The reason can be that they are crystalline - if so, the orientation of the crystal lettice of (one, a few, or) many individual crystal grains has an influence on the properties of the compound material. The individual crystal lettice orientations can be at random or their can be a preferred orientation.
A whole branch of mineralogy and material science deals with the analysis of materials which show crystal preferred orientations (CPOs).
The so-called Orientation Distribution Density Function (ODF) describes how the crystal axes of mineral grains in a sample are oriented relative to an outer coordinate system (e.g. geographic XYZ coordinates). From the ODF, usely depicted as a couple of stereographic density plots (e.g. plots for the crystal a-axis, b-axis and c-axis orientations) the direction and sense of shearing of a rock can be inferred.
Crystal orientations in mineralized tissue are, of course, not defined relative to a geographic coordinate system but to the anatomic directions (or the axis of accretionary growth) of the animal as a reference system.
Coming from the tectonical side I was participating in a workshop on "Textures & Microstructures in Geosciences" in 2005: I was really surprised that there is an application of texture analysis in palaeontology and that some people are really doing it.
U-stage and EBSD are involving single grain measurements of a thin section: For each grain in a certain raster the individual crystal orientation is determined - either by its optical properties using the universal stage (this old-fashioned manual method is rather time-consuming) or by the way how electrons from an electron microbeam are backscattered on a detection screen (Electron Backscatter Diffraction, EBSD). The detected Kikuchi line patterns are indicative for how an individual crystal is oriented (they can be interpreted automatically). What you get in both approaches is not only an ODF but also a crystal orientation map of your thin section.
XRD, Neutron Diffraction. In these methods an X-ray or neutron beam is used to measure a larger volume of a sample comprising several crystals, thereby neutron radiation can penetrate even larger samples completely while the X-ray has a relatively low depth of penetration. From the detected line sprectra pole figures, representing the distributions of certain crystal lettice plane orientations can be derived (from which in turn the ODF of all the crystals in the measured sample volume can be deduced).
The authors employ X-ray diffraction measurements and demonstrate thate the microstructures and crystallographic textures of aragonite layers of species from different mollusc taxa including bivalves, cephalopods, gastropods and monoplacophorans are highly specific and contain a phylogenetic signal: closer relatives are more similar in their shell's crystal orientations.
Pyzalla, A.R., Sander, P.M., Hansen, A., Ferreyo, R., Yi, S.-B., Stempniewicz, M. & Brokmeier, H.-G. (2006): Texture analysis of Sauropod bones from Tendaguru. - Material Science and Engineering A 437: 2-9.
This neutron diffraction approach addresses the question whether the apatite crystallite textures in adolescent and adult Brachiosaurus long bones show some signal indicative for specialized crystal orientations which can be attributed to the giant growth of sauropod dinosaurs. However, comparing their results to the measurements of turkey and other dinosaur long bones they found no significant difference in texture strength or in the predominant direction of fibres.
The reason can be that they are crystalline - if so, the orientation of the crystal lettice of (one, a few, or) many individual crystal grains has an influence on the properties of the compound material. The individual crystal lettice orientations can be at random or their can be a preferred orientation.
A whole branch of mineralogy and material science deals with the analysis of materials which show crystal preferred orientations (CPOs).
From Tectonics...
Structural geologists, who are dealing with microscopic phenomena of tectonic deformation, use mineral textures of rocks, i.e. crystal preferred orientations of the rockforming minerals, as indicators of tectonic movement and deformational regimes (temperature, pressure conditions; deformation rate).The so-called Orientation Distribution Density Function (ODF) describes how the crystal axes of mineral grains in a sample are oriented relative to an outer coordinate system (e.g. geographic XYZ coordinates). From the ODF, usely depicted as a couple of stereographic density plots (e.g. plots for the crystal a-axis, b-axis and c-axis orientations) the direction and sense of shearing of a rock can be inferred.
...to Paleontology
Not only tectonic forces are governing the crystalline properties of natural materials. In a similiar way the (often monomineralic) mineralized tissues of organisms are underlying specific biological formation conditions - the different layers of a skeleton can show rather perfect CPOs.Crystal orientations in mineralized tissue are, of course, not defined relative to a geographic coordinate system but to the anatomic directions (or the axis of accretionary growth) of the animal as a reference system.
Coming from the tectonical side I was participating in a workshop on "Textures & Microstructures in Geosciences" in 2005: I was really surprised that there is an application of texture analysis in palaeontology and that some people are really doing it.
Methodology
You can do either single grain measurements or methods integrating all orientations of crystal grains in a certain volume of the sample.U-stage and EBSD are involving single grain measurements of a thin section: For each grain in a certain raster the individual crystal orientation is determined - either by its optical properties using the universal stage (this old-fashioned manual method is rather time-consuming) or by the way how electrons from an electron microbeam are backscattered on a detection screen (Electron Backscatter Diffraction, EBSD). The detected Kikuchi line patterns are indicative for how an individual crystal is oriented (they can be interpreted automatically). What you get in both approaches is not only an ODF but also a crystal orientation map of your thin section.
XRD, Neutron Diffraction. In these methods an X-ray or neutron beam is used to measure a larger volume of a sample comprising several crystals, thereby neutron radiation can penetrate even larger samples completely while the X-ray has a relatively low depth of penetration. From the detected line sprectra pole figures, representing the distributions of certain crystal lettice plane orientations can be derived (from which in turn the ODF of all the crystals in the measured sample volume can be deduced).
Examples from Paleontology
Chateigner, D., Hedegaard, C. & Wenk, H.-R. (2000): Mollusc shell microstructures and crystallographic textures. - Journal of Structural Geology 22: 1723-1735.The authors employ X-ray diffraction measurements and demonstrate thate the microstructures and crystallographic textures of aragonite layers of species from different mollusc taxa including bivalves, cephalopods, gastropods and monoplacophorans are highly specific and contain a phylogenetic signal: closer relatives are more similar in their shell's crystal orientations.
Pyzalla, A.R., Sander, P.M., Hansen, A., Ferreyo, R., Yi, S.-B., Stempniewicz, M. & Brokmeier, H.-G. (2006): Texture analysis of Sauropod bones from Tendaguru. - Material Science and Engineering A 437: 2-9.
This neutron diffraction approach addresses the question whether the apatite crystallite textures in adolescent and adult Brachiosaurus long bones show some signal indicative for specialized crystal orientations which can be attributed to the giant growth of sauropod dinosaurs. However, comparing their results to the measurements of turkey and other dinosaur long bones they found no significant difference in texture strength or in the predominant direction of fibres.
Prospects
Given the elaborateness of most approaches, measurements of crystallographic textures are rarely used in paleontology - I suppose this will change if it turns out that the analysis of the crystal orientations can provide substantial information which is not obtained from the usual analysis of skeletal histology and microstructures.Donnerstag, 1. Januar 2009
Happy Darwin Year!
Coming to us in 2009: festivities and colloquia celebrating the 200th birthday of Charles Darwin and the 150th anniversary of his grand oeuvre
On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life,
which was "introducing sense into biology" (badly corrupted from the Ernst Mayr quote).
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Looking back a palaeoherpetologist would call 2008 the year of the turtle:
With the descriptions of Odontochelys and Chinlechelys and some further contributions (a list is provided by N. Gardner) the origins of turtles were greatly enlightened.
This topic was variously covered by the pal(a)eo and evo blogs – see for example here and here (Bill Parker's Chinleana) or here and here (Adam Yates' Dracovenator blog).
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BTW: How many recent On-papers do you know?
What is the right occasion to start a contribution with 'on'? Did you ever think about it ... to me such a beginning suggests a long-winded reflection of/ a comment on sth. rather than focusing on the presentation of new data.
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The Twelve Days of (Paleo) Christmas are not yet over - you are still welcome to participate in the poll. >>>>>
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I support Scientific Triassicism! An initiative by Neil from Microecos. What do you think: Is 'Triassic Science' or 'Triassology' a sexier term?
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(How long one week and a half can be: I forgot how to compose a text, that's why this post is only a list of notes.)
On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life,
which was "introducing sense into biology" (badly corrupted from the Ernst Mayr quote).
-----
Looking back a palaeoherpetologist would call 2008 the year of the turtle:
With the descriptions of Odontochelys and Chinlechelys and some further contributions (a list is provided by N. Gardner) the origins of turtles were greatly enlightened.
This topic was variously covered by the pal(a)eo and evo blogs – see for example here and here (Bill Parker's Chinleana) or here and here (Adam Yates' Dracovenator blog).
-----
BTW: How many recent On-papers do you know?
What is the right occasion to start a contribution with 'on'? Did you ever think about it ... to me such a beginning suggests a long-winded reflection of/ a comment on sth. rather than focusing on the presentation of new data.
-----
The Twelve Days of (Paleo) Christmas are not yet over - you are still welcome to participate in the poll. >>>>>
-----
I support Scientific Triassicism! An initiative by Neil from Microecos. What do you think: Is 'Triassic Science' or 'Triassology' a sexier term?
-----
(How long one week and a half can be: I forgot how to compose a text, that's why this post is only a list of notes.)
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