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Nuevos ejemplares del Oligoceno superior de los sedimentos lacustres de la dolina Oberleichtersbach (Baja Franconia, Baviera, Alemania).

RESUMEN: En este articulo se estudian nuevos ejemplares del Oligoceno superior de los sedimentos lacustres de la dolina Oberleichtersbach (Baja Franconia, Baviera, Alemania), determinados como de los galapagos Palaeoemys bessiaca Schleich, 1994 y Palaeomauremys tuberculata (Portis, 1882), las tortugas de caparazon blando Allaeochelysparayrei Noulet, 1867 y cf. Trionyx triunguis Forskal de 1775, y la tortuga mordedora Chelydrasia decheni (V. H. Meyer, 1852). Se discuten las caracteristicas morfologicas de estas cinco especies, su posicion taxonomica y sus implicaciones paleobiologicas. El nuevo material de Palaeomauremys sugiere que P. mlynarskii (Hervet y Lapparent de Broin, 2000) es un sinonimo de P. tuberculata (Portis, 1882).

Palabras clave: Testudines, Palaeoemys bessiaca Schleich, 1994, Palaeomauremys tuberculata (Portis, 1882), Allaeochelys parayrei Noulet, 1867, Trionyx cf. triunguis Forskal, 1775, Chelydrasia decheni (H. V. Meyer, 1852), Oligoceno superior, Chattiense, Oberleichtersbach, Baja Franconia (Alemania), descripcion, paleobiologia.

ABSTRACT: New Late Oligocene (Chattian) remains of the terrapins Palaeoemys hessiaca Schleich, 1994 and Palaeomauremys tuberculata (Portis, 1882), the soft-shelled turtles Allaeochelys parayrei Noulet, 1867 and Trionyx cf. triunguis Forskal, 1775, and the snapping turtle Chelydrasia decheni (H. V. Meyer, 1852) from lacustrine sediments of the Oberleichtersbach doline (Lower Franconia, Bavaria, Germany) are reported. The morphological features of these five species, their taxonomic position and their palaeobiological implications are discussed. The new Palaeomauremys material suggests that P. mlynarskii (Hervet & Lapparent de Broin, 2000) is a junior synonym of P. tuberculata (Portis, 1882).

Key words: Testudines, Palaeoemys hessiaca Schleich, 1994, Palaeomauremys tuberculata (Portis, 1882), Allaeocbelys parayrei Noulet, 1867, Trionyx cf. triunguis Forskal, 1775, Chelydrasta decbeni (H. V. Meyer, 1852), Late Oligocene, Chattian, Obedeichtersbach, Lower Franconia, description, palaeobiology.



From the German terrestrial and limnic deposits of Late Oligocene (Chattian) age the terrapin turtle Palaeomauremys mlynarskii (Hervet & Lapparent de Broin, 2000) was already described from Rott (North Rhine-Westphalia), Enspel (Westerwald: Rhineland-Palatinate), and Oberleichteisbach (Bavaria: Lower Franconia). In the meantime, additional material is available from the Walter Heck collection (Oberleichtersbach) which enables the revision of Emys tuberculata Portis, 1882, a species of previously uncertain status which was assigned to Palaeomauremys Hervet, 2004 by HERVET (2004) and KARL & WETTLAUFER (2010). Based upon the now known morphological features in the new specimens, both taxa can easily be synonymized on generic level.

The genesis and topography of the Late Oligocene (Chattian) doline filling of Oberleichtersbach near Bad Bruckenau (fig. 1) is discussed in detail by MARTINI (2008) which contribution is explicitely referred to. In the same volume BOI-IME (2008) presented a complilation of 43 taxa of fish, amphibians and reptiles. The first consist of six species in two families: five species of Cyprinidae and the new Cobites primigenus Bohme, 2008 (Cobitidae). The amphibians include eight species of Allocaudata and Urodela as well as six species of Anura. The reptiles are present with one species of Diplocynodon (Crocodylia) and 15 species of Squamata: Lacertilia, Anguimorpha, and Ophidia, the latter with the new Texasophis hecki Bohme, 2008. Many of these species present the most ancient record of their groups, one of them is the latest one. With this content of fossils, this locality shows the highest diversity of ectothermic vertebrates of the Late Oligocene world-wide. Additionally there are six species of Testudines. One pleural of "Palaeomauremys mlynarskif' is figured (pl. 4, fig. 7a-b), the other taxa are listed without figures and catalogue numbers. Here it is referred to under the discussion of the species in question.

This paper is supplement 180 of chapter "C. Fourth faunal change: Paleogene-Neogene (P/N) boundary/V. Rise of stream terrapins", in: Studia Palaeocheloniologica, IV (KARL, 2011).



Ordo Testudines Linnaeus, 1758

Suprafamilia Testudinoidea Batsch, 1788

Familia Geoemydidae Theobald, 1868 (= Bataguridae Gray, 1869)

Subfamilia Palaeochelyinae Karl & Wettlaufer, 2011

Genus Palaeoemys Schleich, 1994

--1994 Palaeoemys n. g. Schleich: 82-87.

--2004 Palaeoemys Schleich, 1994--HERVET: 65-70.

--2004 Palaeoemys Schleich, 1994--CLAUDE & TONG: 6.

--2011 Palaeoemys Schleich, 1994--KARL & WETTLAUFER: 179.

Type species: Palaeoemys hessiaca Schleich, 1994: 82.

Additional species: Palaeoemys occitana Hervet, 2004, Palaeoemys testudiniformis (Owen, 1842) emend. CLAUDE & TONG (2004).

Geographic and stratigraphic distribution:

Early Eocene (MP9 to MP 10 boundary), Saint-Papoul near Castelnaudary (Aude, France) and Monthelon near Epernay (Marne, France: MP 19). Middle Eocene (MP11 to MP 13), Borken (northern Hesse, Germany).

Original diagnosis: SCHLEICH (1994: 82). With the present specimen the stratigraphic distribution is extended to the Late Oligocene and thus to the late Palaeogene.

3.1.1. Palaeoemys cf. hessiaca Schleich, 1994

--2008 Emydidae indet. Bohme: 168.

New material: OLB 2-1, rigth epiplastron in ventral view (plate 4, figs. 1-2); OLB 2-2, rigth pleural I remain (plate 4, figs. 3-4); OLB 2-3, pleural remain (plate 4, figs. 5-6); OLB 4-1, phalanx III (plate 4, fig. 7); OLB 4-2, scapula remain (plate 4, fig. 8); OLB 2-4, left hyoplastron remain (plate 4, figs. 9-10); OLB 2-4, left hyoplastron remain (plate 4, figs. 9-10).

Discussion: The hitherto single record of Palaeoemys. Further specimens possibly belonging to this taxon are too fragmentary for a definite assignment. The ventral and visceral course of the gular suture, the furrow of the skin margin and the extremely week gular bulge exactly resemble the known morphology of this genus (see HERVET, 2004; SCHLEICH, 1994), but due to the preservation the specimen is only tentatively assigned to P. hessiaca.


3 2. Genus Palaeomauremys Hervet, 2004 Selected synonymy:

--1882 Emys--Portis: 19-20.

--2000 Palaeochelys Schleich 1994--HERVET & LAPPARENT DE BROIN: 563.

--2004 Palaeomauremys n. gen. HERVET: 54-56.

--2011 Palaeomauremys Hervet, 2004--KARL & WETTLAUFER: 175.

Type species: Emys titberculata PORTIS, 1882.

Original description: PORTIS (1882: 19-20, pl. 4).

Type locality: Rochette near Lausanne (Switzerland).

Type straaim: Subalpine Molasse (Molasse d'eau douce inferieure, USM), Late Oligocene: MP 28-30 (ENGESSER, 1990), MP 29 (ENGESSER & MODDEN, 1997).

Holotype: MGL 8901 (= PORTIS, 1882, pl. 4), anterior fragment of carapace + fragments of plastron (pl. 76, figs. E, F, G and H).

Emended description by HERVET (2004): "Le repli ventral de la cervicale est de longueur moyenne sur la face interne de la nucale, il est tres large (Pl. 76, fig. G). Le repli ventral des M1 est assez long et il a presence de deux legers bourrelets transversaux sur la face interne de la nucale. La face interne de la PL1 montre l'insertion du pilier axillaire sur environ un tiers de la larger de la pleurale (Pl. 76, fig. H); il y a un leger bourrelet osseux sur la PL1, medialement a cette insertion". For the generic diagnosis of the carapace: see HERVET (2003, 2004), in particular map 4.11 and chapter 4 in the paper in 2004.

Remarks: Most important synonym for the region in question is Palaeochelys mlynarskii (Hervet & Lapparent de Broin, 2000) from the Late Oligocene (MP 30: BOHME & LANG, 1991; MORS, 1995) of the lignites of Rott near Bonn (North Rhine-Westphalia, Germany; see: HERVET & LAPPARENT DE BROIN, 2000; HERVET 2003 and 2004, with generic diagnosis). For the Enspel specimen Exemplar: see KARL & WETTLAUFER (2011). A small species with a carapace length of originally only 125 mm, but the present material suggests a length of more than 200 mm.



3.2.1. Palaeomauremys tuberculata (portis, 18820

Selected synonymy:

--1882 Emys tuberculata n. sp. Portis: 19-20; pl. 4.

--1892 Testudo Riedli nov. form. Hoemes: 243 (synonyms in KARL, 1994/95).

--2000 Palaeochelys mlynarskii n. sp. Hervet & Lapparent de Broin: 563-569; figs. 1-3.

--2004 Palaeomauremys <<tuberculata>> (Portis, 1882)--HERVET: 163.

--2011 Palaeochelys mlynarskii Hervet & Lapparent de Broin, 2000--KARL & WETTLAUFER: 175-177.

--2011 Palaeomauremys "tuberculata" (Portis, 1882)--KARL & WETTLAUFER: 175.

Geographic and stratigraphic distribution: (i) Germany: Top of Late Oligocene (MP 30), Rott near Bonn (North Rhine-Westphalia); Enspel Maar (Rhi-neland-Palatinate: Westerwald); doline filling at Oberleichtersbach (Bavaria: Lower Franconia; KARL & WETTLAUFER, 2011).

(ii) Slovenia: Trbovlje (former Trifail).

New material: OLB 1-1, anterior carapace remain (plate 1, figs. 1-2); OLB 1-2, left pleural II (plate 1, figs. 3-4); OLB 1-3, pygal (plate 1, fig. 5); OLB 1-4, OLB 1-5, OLB 1-6, bridge peripherals (plate 1, figs. 6-8); OLB 1-9, rigth anterior plastral lobe (plate 2, figs. 7-8); OLB 110, rigth epiplastron (plate 2, figs. 1-2); OLB 1-11, left epiplastron (plate 2, figs. 3-4); OLB 112, left xiphiplas-tron; OLB 1-13, rigth xiphiplastron remain (plate 2, figs. 9-10).

Main characters of the new material: Length of carapace = 200 mm. Cervical distinctly developed, visceral part larger than dorsal part; epiplastron strongly vaulted, with distinctly marked geoemydaloid gular bulge; gular furrows cutting entoplastron, humeropectoral furrows not; entoplastron 1,2 x broader than long, rhomb-like, anteriorly markedly pointed; anal notch obtuse-angled; pygal geoemydaloid; neurals emydoidal and geoemydaloid; three (to maybe five) strong dorsal keels, no keel on bridge developed; bridge piers week, inguinal piers inserting into pleural I via peripherals II and III, peripherals III with distinct visceral moschus channel, all marginal furrows situated on peripherals; typical geoemydaloid structure of areols on carapace present, surface of carapace with coarse sculpture, partly with very deep furrows on the horn-scutes. In fragmentary material the latter can feign the presence of terrestrial turtles: "Several plastron and carapace remains belong to an undetermined species of Testudo ..., and a second land turtle known only by one plastron fragment (not figured). It differs from Testudo in having rising (= geochelonid according to SCHLEICH 1988) sutural furrows, typical for large land turtles of the genus Geochelone sensu lato" (BOHME, 2008).

KARL (1994/95) grouped Testudo riedli Hoernes, 1892 with Clemmydopsis Boda, 1927. Based upon the new material from Oberleichtersbach it is almost sure that it really belongs to Palaeomauremys. There are exactly the same morphological features in the anterior part of the internal mold from Trbovlje (former Trifail) in Slovenia and in the anterior visceral region of the carapax from Oberleichtersbach. Even the structtire of the plastron is the same. The close stratigra-phical age of the Slovenian and German specimens support their close relationships (plate 3, figs. 1-2). According to RIJAVEC & DOZET (2002) the limno-brackish Socka Beds (former Sotzkaschichten) are subdivided into the Lower Socka Beds, the main coal horizon, and the Upper Socka Beds. They are overlain by the Late Oligocene Marine Marly Clay, dacito-andesitic tuff and pyroclastic breccia. The stratigraphical position is Egerian (= Late Oligocene: Chattian to the Oligocene/ Miocene boundary beds). For a detailed description and the synonymy of the taxon see KARL (1994/95) and GROSS (1994). The Egerian in the Paratethys largely corresponds with the late Chattian and the complete Aquitanian (25,8-20,3 MY) (BALDI, 1975) and includes the "Gosau-Schichten".

During railway constructions the visceral mold of a turtle carapace was discovered in a quarry for building stones in the "Gosau-Schichten" at the Wuhrberg S Windischgarsten (Upper Austria) and donated to the k. k. Geolo-gischen Reichsanstalt in Vienna by Ing. Gross (Spital, Pahrn, Austria) in 1905. The comparable characters completely correspond with those of the steinkern from Trbovlje (former Trifail, Slovenia). Thus the Wuhrberg specimen has to be assigned to Palaeomauremys, too (plate 3, fig. 3). With a length of 254 mm and a width of 186 mm this carapace is the largest known.


Suprafamilia Trionychoidea Fitzinger, 1826

Familia Carettochelyidae Boulenger, 1887

Subfamilia Carettochelyinae Boulenger, 1887

3.3. Genus Allaeochelys Noulet, 1867

Synonymy: See KUHN (1964), BROIN (1977) and LAPPARENT DE BROIN (2001).

Type species: Allaeochelys parayrei NOULET, 1867 (original generic designation); originally described from Eocene deposits in southern France.

Diagnosis: See MLYNARSKI (1976: 73-74).

Remarks: Neither the genus name Allaeochelys nor its author NOULET (1867) were listed in the "Fossilium Catalogus" on "Trionychia fossilia" (HUMMEL, 1932) (see KARL et at, 2006).

Geographic and stratigraphic distribution: Early to late Middle Eocene in southern France, Spain, England, Belgium and Germany, in Germany up to the Late Oligocene (KARL et al., 2006).

3.3.1. Allaeochelys parayrei Noulet, 1867

Selected synonymy:

--1956 Anosteira crassesculpta Harrassowitz, 1922--GRAMANN: 18.

--1956 Anosteira crassesculpta = gracilis Harrassowitz, 1922--KUHN: 181.

--1986 "Anosteira" aff. crassesculpta Harrassowitz, 1922--SCHLEICH: 285.

--2006 Allaeochelys parayrei Noulet, 1867 (syn. Anosteira crassesculpta et A. gracilis Harrassowitz, 1922)--KARL, GRONING & BRAUCKMANN: 51-57.

--2007 Allaeochelys Noulet, 1867 (syn. Anosteira crassesculpta Harrassowitz, 1922)--KARL, 2007: 61.

New material: Left dental (adult) and right xiphiplastron (juvenile).

Brief description: Sculpture on xiphiplastron flat and reticular, alveolar plain of dental without sculpture (withotit marked chewing ridges). Fossa meckeli open until symphysis. Labial margin with the typical porous structure of Trionychoidea.

Familia Trionychidae Fitzinger, 1826

Subfamilia Trionychinae Fitzinger, 1826

Subtribus Trionychina Fitzinger, 1826

3.4. Genus Trionyx Geoffroy Saint Hilaire, 1809

Synonymy: See KARL (1998), older synonymy in HtlMMEL (1929, 1932), LOVERIDGE & WILLIAMS (1957), WERMUTH & MERTENS (1961), KUHN (1964), and IVERSON (1992).

Type species: Testudo triunguis Forskal, 1775.

Terra typica of type species: "In Nilo rarior" [= Nile River] (IVERSON, 1992).

Diagnosis: See KARL (1998).

Remarks: As pointed out by KARL (1999), the Eocene/Oligocene transition is marked by a change in the Trionychidae in Central Europe. The Eocene Nearctic Rafetoides austriacus (Peters, 1868) disappear and open wide space for the oriental species Trionyxgregarius/triunguis. The latter becomes the dominant species of the Neogene Palaearctic an is now restricted to a postglacial relict areal in Ethiopia flor the history of distribution see KARL, 1999). Trionyx triunguis is also known from the Early Oligocene sequence in the WeiIgelster Basin (Saxonia, Germany; KARL, 2007). On the other hand, in the Early Oligocene "Melanien-Ton" of Borken (Hesse, Germany) Rafetoides austriacus still occurs (KARL & MULER, 2008).

Geographic and stratigraphic distribution:

(i) Late Oligocene to Miocene: Austria, Slovenia, southern and Central Germany, France, Hungary, Romania, Bohemia, Switzerland.

(ii) Miocene to Pliocene: Germany, Egypt.

(iii) Tertiary (without exact data): Nupur in Nepal.

(iv) Prehistoric: Egypt.

(v) Recent: Southern Turkey to Africa (Senegal to Angola, Somalia and Egypt) (IVERSON, 1992).

The additional species Trionyx gregarius (syn. Amyda gregaria Gilmore, 1931) was memioned from the Early and "Middle" Oligocene Houldjin Formation at Camp Margetts, 25 miles SW Iren Babasu, Inner Mongolia (= Neimenggu, northern China).

3.4.1 Trionyx cf. triunguis (Forskal, 1775)

Remarks: According to BOHME (2008) "the rearest turtle in Oberleichtersbach are the soft-shell of river turtle Trionyx sp., documented by one characteristically sculptured pleural fragment (not figured)". Here only a single free pleural cone, showing the structured transition of the proximal part to the pleural plate.

Superfamilia Chelydroidae Gray, 1831

Familia Chelydridae Agassiz, 1857

Subfamilia Chelydropsinae Mynarski, 1980

3.5. Genus Chelydrasia Chkhikvadze, 1999

Synonymy: See KUHN (1964), DE BROIN (1977), DE LAPPARENT DE BROIN (2001) and HUTCHISON in STEYERMARK et al. (2008).

Type species: Chelydropsis minax Chkhikvadze, 1971 (?= Chelydra decheni H. V. Meyer, 1852).

Diagnose: CHKHIKVADZE (1999).

Geographic and stratigraphic distribution: Early Eocene to Pliocene in Kazakhstan; Middle Eocene to Early Miocene in Europe (HUTCHISON in STEYERMARK et al., 2008; KARL et al. in print).

Remarks: CHKHIKVADZE (1999) established Chelydrasia with its original type species Chelydropsis minax Chkhikvadze, 1971 from the Early Oligocene of the Zajsan Basin in Eastern Kazakhstan. He also assigned Ch. sanctihenrici and all the species of the so-called decheni-sanctihenrici group. Furthermore, he assumed an uncertain taxon from Artenay (MN4) to be closely related. According to HUTCHISON in STEYERMARK et al. (2008) Chalydrasia now includes Ch. minax Chkhikvadze, 1971, Ch. poena Chkhikvadze, 1971, Ch. apellanzini (Murelaga et al., 1999), Ch. decheni (H. V. Meyer, 1852), Ch. sanctihenrici (De Broin, 1977), "Ch." kusnetzovi (Chkhikvadze, 1985). Judging from the preservation, the differences between Ch. minax and Ch. decheni seem to be not definitely clarified. If Ch. minax is regarded a junior synonym of Ch. decheni the latter becomes the type species.

3.5.1. Chelydrasia decheni (H. V. Meyer, 1852)

Synonymy: See KUHN (1964), DE BROIN (1977), DE LAPPARENT DE BROIN (2001), KARL et al. (in print).

New material: OLB 3-1, left pleural I in dorsal view in situ at grey chalky sediment (plate 4, fig. 11); OLB 3-2, pleural remains in dorsal view in situ at reddish chalky sediment with many gastropods (plate 4, fig. 12).

Brief description: The dimensions of pleural I are superficially similar to those of Palaeomauremys, but with a thickness with no more than 1 mm in its proximal region it is distinctly thinner. The plate thickens towards the margin but does not even approximately reaches the dimensions of the following one. The dorsal plain is smooth, with a few areoles only towards the lateral margin, but without any rudiments of keels. The latter first appear on a few smaller fragments of pleurals.




The palaeobiological conditions of the extinct genus Palaeomauremys are supposed to be very similar to those of the Recent and fossil Mauremys. Today these Palaeochelyinae inhabit calm smaller water systems as pools, ditches, of muddy brooks from the lowland up to the mountainous regions. Recent species are omnivorous (see for example KARL & WETTLAUFER, 2011).

The most detailed climatic data, based upon plant remains, were presented by Mm (2008). The annual average temperature varied between 10 and 16[degrees]C, the warmest month reached about 22 to 28[degrees]C whereas the temperature of the coldest month changed from about -3 to -8[degrees]C. The annual precipitation ranged presumably from 630 to 1040 mm, with one or two dry months of only 18 to 7 mm.

Comparable Recent conditions are known only from a few Chinese weather stations, as for example Enshi (= Engshih) in Hubei (= Hupeh) as well as Tai Shan and Jinan (= Tsinan) in Shandong (= Schantung). For the time interval between 23.5 and 25 million years ago MAl (2008) proposed a new and separate "floral complex Oberleichtersbach" which is a particular Carpino-Ostryon (MMF) assemblage including semihumid species, typical floral elements of dry summers and a great number of annual plants of open places.

According to BOHME (2008), the fauna suggests the existence of a large, shallow, permanent and oxygen-rich lake with intensively structured shore and an afflux. The faunal climatic conditions were considered to be subtropical and humid which is not completely in accordance with the palaeobotanical data.


In contrast to other authors, CLAUDE & TONG (2004) synonymized a large number of generic names of the Palaeochelyinae with Palaeoemys which is regarded as a taxon different from Mauremys. In particular they assign Palaeoemys to the so-called Malayemys complex, including Malayemys, Geoclemys, Palaeoemys, and ?Borkenia. On the other hand, Mauremys is allocated to the Melanochelys complex, including Melanochelys, Mauremys, Sacalia, Cuora, Cistoclemmys, Notochelys, Heosemys, Hieremys, Cyclemys, Leucocephalon, Ocadia, Chinemys, Palaeochelys, ?Siebenrockiella and ?Geoemyda. These different views implicate a hotspot on the European spit during the Paleogene as it was already pointed out by KARL (1998, 1999) and DANILOV (2005) for the Trionychidae. Likewise this seems to be true for the Palaeochelyinae. Within the whole relationship distinct tendencies can be recognized for splitting into the genenc complex Mauremys/Ocadia, the Ptychogastrinae and the Geoemydinae, still combined with large overlapping (see for example KHOZATSKY & MLYNARSKI, 1966). Surely this problem cannot yet finally dissolved since BROPHY et al. (2006) show twelve cases of intergeneric hybridisation, where the following genera are connected by genetic overlapping: Mauremys x Chinemys, Mauremys x Cuora, Mauremys x Cyclemys, Mauremys x Heosemys, Mauremys x Sacalia, Cuora x Geoemyda und Cuora x Sacalia, i.e. the complete complex with its fossil roots in the Palaearctic. Indeed, even recently often different "hybrid species" occur in open nature, as for example Cbinemys megacepbala, Mauremys pritchardi, Ocadia glyphistoma, Ocadia philippeni, Mauremys iversoni and others. The latter is even breeded in Chinese turtle farms as a hybrid of Mauremys mutica x Cuora trifasciata and was already discussed under taxonomic respect (FONG & CHEN, 2010; PARHAM & CHI, 2001). Consequently, FRITZ & HAVAe (2007) regarded Chinemys, Sacalia und Ocadia etc. as synonyms of Mauremys as a conclusive result of a genetic analysis of single individuals, just in contrast to the comparison of morphologically studied series. Once more, this clearly shows that precipitate "results" based only on genetic analyses are not sufficient as they are not compatible with morphological conclusions.


Unser herzlicher Dank gilt Frau Dr. Irene Zom, Geologische Bundesanstalt Wien, Dr. Martin Gross, Steiermarkisches Landesmuseum Joanneum Graz und Herm Walter Heck, Oberleichtersbach fur die freundliche Bereitstellung des Materials aus ihren Sammlungen.


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Hans-Volker KARL (*,**)

Elke GRONING (***)

Carsten BRAUCKMANN (***)

(*) Thuringisches Landesamt fur Denkmalpflege und Archaologie. Humboldtstrasse 11. D-99423 Weimar. Correo-e:

(**) Geoscience Centre of the University of Gottingen. Department Geobiology. Goldschmidtstrasse 3, D-37077 Gottingen.

(***) Institut fur Geologie und Palaontologie. TU Clausthal. Leibnizstrasse 10. D-38678 Clausthal-Zellerfeld. Germany. Correo-e:;

(FECHA DE RECEPCION: 2012-01-26) (FECHA DE ADMISION: 2012-01-30) BIBLID [0211-8327 (2011) 47 (2); 175-194]
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Author:Karl, Hans-Volker; Groning, Elke; Brauckmann, Carsten
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Date:Jul 1, 2011
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