Chroniosuchia: Paper on osteoderm histology in online preview

...my first experience with bone histology:

Buchwitz, M., Witzmann, F., Voigt, S. & Golubev, V. in press. Osteoderm microstructure indicates the presence of a crocodylian-like trunk bracing system in a group of armoured basal tetrapods. Acta Zoologica, DOI: 10.1111/j.1463-6395.2011.00502.x

Abstract. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel-fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel-fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio- or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian-like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.

Maths in Paleontology (I): Data

''In every special doctrine of nature only so much science proper can be found as there is mathematics in it.'' - Immanuel Kant, Metaphysical Foundations of Natural Science (1786)

Warningly the maths professor who got the unthankful task to teach us first-semester scientists-to-be some basic basics of his field chose Kant's statement as the first in his first lecture on "higher" maths. However, when I started my studies in geology and paleontology, there was another saying among old school geology teachers: "A bad mathematician makes a good geologist."

Many a fellow student were rather willing to believe in these latter words than in the inconvenient alternative. (I always considered this believe as outdated and I got the feeling that geology as a science might have been shaped not only by the talents of its protagonists but also by their limitations in terms of exactness and rigorousity.)

Luckily you were not necessarily considered as a bad geologist if you were interested in maths and the notion that modern geoscience involves maths and exact methods (e.g. methods of quantitative data analysis, databases, multivariate statistics and geostatistics, geoinformatics and geographic information systems, 3D and 4D modelling, remote sensing) was clearly on the rise. Perhaps from a biologists' point of view this story would be different, but, to tell you the truth, some of the biology-based paleontologists I got to know are not much living on the exact side either.

Apart from microscopy seminars, field, and lab practicals which teach you ways of data acquisition some classes in statistics and data analysis during first semesters of study give you an idea about the structure of data and ways how to sample and how to deal with data in order to find new knowledge, e.g. a relationship between two phenomena previously not considered to be related.

At the very beginning you will learn that there are different types of data used in paleontology and that you have to bring your data into shape for any kind of mathematical analysis tools, i.e. arrange them as a data table such as the following:

Specimen	Class	State of XYZ	No. of UVW	size L [mm]	size M [cm²]
A	a	Aa	2	12.1	234
B	b	B	3	13.3	87
...	...	...	...	...	...
X	x	X	...	...	...

Normally lines of the table represent samples (or groups of samples or taxa) whereas columns may represent various features or measures. Such features may be the belonging to a certain class or category or the presence, absence, or specificity of a feature. Measured values as entries may have a discrete contribution (e.g. natural numbers such as the number of teeth or segments or body chambers) or a continuous distribution (e.g. length, area, angle, temperature measurements).

Various data relevant for paleontologists can be arranged as tables, such as morphological and microstructural data, stable isotope and other geochemical data, geographical, sedimentological, and stratigraphic data, as well as taphonomic and paleoecological data. Some of these data have a special structure and can be referred to one of the following types:


Compositional data...

... add up to 100%. Chemical compositions of fossils or faunal compositions are compositional data:

Community	Trilobites	Brachiopods	Echinoderms	Poriferans	Nautiloids
A	23 [%]	42	17	5	13
B	10	15	5	50	20
...	...	...	...	...	...
X	...	...	...	...	...

These data require careful considerations and a special kind of maths because all variables are (necessarily) correlated and thus an alleged dependence, e.g. of brachiopod and echinoderm abundances, can be obscured by variation in another group.


Spatially or temporally correlated data

‘Spatial correlation’ means that values for data points close to each other are more similar than values of more distant data points – e.g. the faunal composition of an ecosystem from Arizona is rather like that of a Nevada community than that of a Massachusetts community.

Locality	Easting (X)	Northing (Y)	Facies	Archosaurs [%]	Rhynchosaurs [%]
A	5687	0487	lacustrine	23	45
B	6485	0808	fluviatile	34	38
C	6800	1490	fluviatile	40	37
...	...	...	...	...	...

Geostatistics is the usual method to deal with spatially correlated data. Spatial correlation can also occur on much smaller scales, e. g. the shape and size of two skull bones in contact to each other can show a stronger dependence than the shape and size of bones that are more distant to each other.

In paleontology temporal correlation is quite abundant, especially if your study considers different stratigraphic ages or sedimentological field data:

Population	Horizon	Ar/Ar age [Ma]	Facies	δ18O [‰]	Average size [mm]
A	1	210 ± 1	deltaic	-2.0	5.2
B	2a	N/A	distal shelf	1.4	6.4
C	2c	207 ± 2	?	2.1	6.8
D	4	200 ± 1	deltaic	-2.2	6.0

As in stockmarket analytics methods of time series analysis can be applied to interpret temporally correlated data (i.e. time series). Such data may be relevant for your study as they often indicate evolutionary trends (biological evolution in the stricter sense but also evolution of paleoenvironments), cyclic processes with a certain periodicity, and/or they can form the basis for relating contemporaneous processes in the geological past (e.g. stratigraphic correlation of separate sedimentary successions).


Orientation data

For elongated fossils such as conical shells or long bones the orientation of the fossil long axis towards the geographical cordinate system can be measured using a compass (with inclinometer). In a similar way the orientation of bedding planes can be documented. Such measurements are often used for the purpose of deducing the former transport direction of a ancient sediment transport and depostion system (such as a river, delta, or alluvial fan). A data table with orientation data may look like that:

Specimen No.	Description	Length [cm]	Horizon	Azimuth>	Dip
1	long bone	21	1	N 20° E	0°
2	rib	12	1	N 10° W	5°
3	calamite stem	80	2a	N 15° E	0°
...	...	...	...	...	...

“Azimuth” refers to the angle towards north. Orientation data are distributed on a halfsphere. Mean values (e.g. the average orientation of long bones) and other distribution parameters cannot be derived directly from the averaging of orientation angles but vector arithmetics has to be applied.


Cladistic data

Phylogeny on the basis of morphology conventionally involves cladistic methods, especially in the field of vertebrate paleontology which deals with a particular character-rich group that is deemed suitable for cladistic approaches employing certain kinds of analysis software specialized for the calculation of phylogenetic trees (e.g. PAUP, WinClada).

In cladistic datasets lines represent taxa, mostly species or genera of the group of interest, and columns represent characters (ordered by number), i. e. features of the skeleton which are variable among the included taxa:

Taxon	1	2	3	4	5	6	7	8	9	10
A-saurus	0	0	0	0	0	0	1	0	?	0
B-raptor	?	0	1	?	0	0	1	1	0	0
C-onyx	1	1	0	?	-	1	-	1	0	1
D-ops	1	1	2	1	1	1	1	0	1	1
E-mimus	0	-	2	1	1	2	0	0	1	1

One of the main issues in cladistics is the definition of characters and the correct (unbiased) coding of morphological information. You can include qualitative differences ("bone X contacts bone Y but not bone Z" = character state “0”; "bone X contacts bones Y and Z" = character state “1”) and quantitative differences ("length of metatarsal 3 larger than or as large as length of metatarsal 4" = character state "0"; "mt3 is shorter than mt4" = "1"). Sometimes mixed character states like "0 or 1 [but not 2]" occur in a taxon and are coded accordingly.


Missing data...

...occur all the time in paleontology ... either because specimens are not complete enough or because their geological age cannot be exactly determined or because specimens are too rare or valuable to use them for a destructive analysis method or because they are for some reason no longer accessible. "N/A" ("not applicable") or empty entries or question marks often symbolize missing data.


Some introductory literature:

Borradaile, G. J. 2003. Statistics of Earth Science Data. Springer, Berlin, 280 pages. ISBN 3540436030

Swan, A. R. H. and M. Sandilands. 1995. Introduction to geological data analysis. Blackwell, Oxford, 446 pages. ISBN 0632032243

New paper on cycadophytes from Madygen

Moisan, P., S. Voigt, C. Pott, M. Buchwitz, J. Schneider, and H. Kerp. in press. Cycadalean and bennettitalean foliage from the Triassic Madygen Lagerstätte (SW Kyrgyzstan, Central Asia). Review of Palaeobotany and Palynology. [DOI:10.1016/j.revpalbo.2010.11.008]

Philippe Moisan who is doing his Ph.D. in Münster (with paleobotanist Hans Kerp as his supervisor) studies the flora of the Triassic Madygen Fm. In his first paper on that issue he introduces cycadophyte finds collected between 2005 and 2009.

Many of the studied the specimen come from the same succession and locality as Madygenerpeton (there is also a small sketch of the sedimentary profile, see Fig. 2).

One thing I learned from this study was that so-called "xeromorphic features", i.e. plant features that are usually the consequence of an adaptation to aridity, cannot only occur in xerophytes, i.e. in plants adapted to dry environments, but (for other reasons) in hygrophytic and halophytic plants as well.

Indications for aridity, such as desiccation crack horizons or or seasonally drying-out ponds and rivers or wide-spread red bed sediments are lacking in Madygen. Thus, according to Philippe's interpretation, "xeromorphism" in Madygen plants probably served other purposes than the xeromorphism of xerophytes (e.g. "self-cleaning of the leaf surface, regulation of excessive radiation and leaf temperature, mechanical defense against phytophagous insects").

Three recent papers on chroniosuchians

Buchwitz M, Voigt S. 2010. Peculiar carapace structure of a Triassic chroniosuchian implies evolutionary shift in trunk flexibility. Journal of Vertebrate Paleontology 30: 1697-1708. [Link]

Schoch RR, Voigt S, Buchwitz M. 2010. A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships. Zoological Journal of the Linnaean Society 160: 515-530. [Link]

Klembara J, Clack J, Čerňanský A. 2010. The anatomy of palate of Chroniosaurus dongusensis (Chroniosuchia, Chroniosuchidae) from the Upper Permian of Russia. Palaeontology 53: 1147-1153. [Link]

The redescription of the Chroniosaurus dongusensis palate by Klembara and colleagues adds further data to the morphological dataset provided by Clack and Klembara (2009) in their revision of C. dongusensis on the basis of a new specimen (which is the most complete of any yet known chroniosuchian). According to the updated phylogenetic analysis from the 2010 paper Chroniosaurus as the only included chroniosuchian taxon formed the sister group of embolomeres.

Schoch and colleagues (me included) describe Madygenerpeton pustulatus, a new species of chroniosuchians from the Middle to Late Triassic of Central Asia with a highly derived skull morphology and a carapace that was chroniosuchid-like in many aspects. The find shows that one lineage of chroniosuchids survived the Permian-Triassic boundary (by 20 or so million years).

The authors discuss characteristics uniting chroniosuchians with "higher reptiliomorphs" and unlike the approach of Klembara and colleagues their cladistic analysis, which includes five chroniosuchian taxa, results in a position of chroniosuchians somewhat closer to amniotes than to embolomeres. Chroniosaurus comes out as the closest relative of Madygenerpeton (both share the characteristic ornamentation of the skull and osteoderms besides other features).

Buchwitz & Voigt consider the functionality of chroniosuchian carapaces, comparing them to archosaur osteoderm systems. They argue that chroniosuchian carapaces basically served terrestrial locomotion but that the higher lateral flexibility of the Madygenerpeton osteoderm system was linked to a secondary increase in undulation swimming capability.

Reference:
Clack JA, Klembara J. 2009. An articulated specimen of Chroniosaurus dongusensis, and the morphology and relationships of the chroniosuchids. Special Papers in Palaeontology 81: 15-42. [Link]

Palges Meeting October 2010 in Munich

The 80 th Annual Meeting of the German Paleontological Society took place from the 6th through the 8th October 2010 within the halls of the Bavarian State Collection for Geology and Paleontology in Munich.

Of particular interest for me was the session on Early Mesozoic vertebrates chaired by the Rauhut couple and Richard Butler as it united many interesting characters, such as Silvio Renesto, Martin Ezcurra, Rainer Schoch, and Daniela Schwarz-Wings and covered a variety of Triassic vertebrates including archosaurs, temnospondyls, and bony fish.

With 5 talks and 6 posters our small Freibergian working group had quite a number of contributions this year (my prof Jörg Schneider was talking about Paleozoic cockroaches from China, Olaf Elicki about Cambrian trace fossils from Africa and the Middle East, Frederik Spindler about the evolution of haptodonts and other early synapsids, Jan Fischer about oxygen isotope signals in Permian and Triassic freshwater shark teeth and I had a talk on osteoderm histology and the Chroniosuchia). My colleagues Ilja Kogan and Jan Fischer won the 1st poster prize with their poster entitled "The Madygen lake deposits: A unique multi-taxa kindergarten for Triassic fisches" - which is quite an achievement as normally the winner comes from the host institute of the Palges Meeting.

The image on the right shows me in front of a poster entitled "Paleontology in the German Wikipedia" [pdf].

Even though there are many private collectors and paleontology enthusisasts in Germany, Austria, and Switzerland you won't find much about "regional paleontology" in the German Wikipedia which was the reason for my colleagues and me to introduce some aspects of Wikipedian (Pop-)Sciencewriting.

The poster praised the advantages Wikipedia can have if it is reasonably incorporated in public outreach campaigns and we commented critically on the dinosaur focus which increases the already biased public image of what paleontology is about.