Morphological fluctuations of ammonoid assemblages from the Muschelkalk (Middle Triassic) of the Germanic Basin—indicators of their ecology, extinctions, and immigrations

Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7 – 34 www.elsevier.com/locate/palaeo

Morphological f luctuations of ammonoid assemblages from the Muschelkalk (Middle Triassic) of the Germanic Basin—indicators of their ecology, extinctions, and immigrations Christian Kluga,T, Wolfgang Schatza, Dieter Kornb, Achim G. Reisdorf c a

Pala¨ontologisches Institut und Museum der Universita¨t Zu¨rich, Karl-Schmidt-Str. 4, 8006 Zu¨rich, Switzerland b Museum fu¨r Naturkunde, Humboldt-Universita¨t zu Berlin, Invalidenstr. 43, 10115 Berlin, Germany c Geologisch-Pala¨ontologisches Institut, Universita¨t Basel, Bernoullistrasse 32, 4056 Basel, Switzerland Received 23 March 2004; received in revised form 14 December 2004; accepted 4 February 2005

Abstract For a stratophenetic analysis of Middle Triassic ammonoids from the German Muschelkalk (Anisian, Ladinian), whorl expansion rates, whorl width indexes, umbilical width indexes, maximal conch sizes, body chamber lengths, the orientations of the aperture, and a number of sculpture parameters of approximately 500 specimens were identified. 274 of these data sets, sorted according to their stratigraphic age, were evaluated in scatter plots as well as canonical discriminant function analyses. Several of the diagrams that were produced in these analyses reflect more or less steady changes in conch morphology through geological time, except for some intervals with abrupt and rather drastic transformations. These morphological discontinuities are synchronous with immigrations into the Germanic Basin of crinoid and brachiopod taxa. This discovery indicates disturbances in the endemic evolution of the ammonoids caused by such immigrations. At a small scale, this case study demonstrates that a rising sea level may have boosted the faunal exchange between an open marine and a restricted epicontinental basin, causing a minor regional increase in biodiversity. It also demonstrates that the evolution of dGermanicT ceratites happened mainly within the Germanic Basin but partially probably also within the Tethyan Sea. The ecology of the ceratites from this Basin is discussed. Accordingly, they were stenohaline, good backward swimmers with a good manoeuvrability, and able to achieve neutral buoyancy. D 2005 Elsevier B.V. All rights reserved. Keywords: Ammonoidea; Triassic; Immigrations; Diversity; Stratophenetic analysis; Morphospace; Ecology

1. Introduction

T Corresponding author. E-mail address: [email protected] (C. Klug). 0031-0182/$ - see front matter D 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2005.02.002

Cephalopod remains are rather abundant in the sedimentary rocks of the Muschelkalk (Anisian and Ladinian, Middle Triassic) of central Europe. In

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contrast to many other epicontinental marine basins of Mesozoic age, specimens of only a few cephalopod genera have been recorded. Among these, specimens of the ammonoid genera Ceratites (the heraldic animal of this journal) and Paraceratites as well as of the nautiloid genus Germanonautilus by far outnumber representatives of other genera with respect to diversity, morphological disparity, and abundance. During the late Anisian and Ladinian, the species of Ceratites and Paraceratites underwent continuous morphological transformation. This transformation is reflected in a fairly large number of species and subspecies as well as in successive biozones (e.g. Philippi, 1898, 1901; Riedel, 1916; Penndorf, 1951; Wenger, 1957; Urlichs and Mundlos, 1987; Urlichs and Vath, 1990; Urlichs, 1993). Because of the putatively simple phylogenetic patterns within Ceratites and Paraceratites (e.g. Mu¨ller, 1954; Wenger, 1957; Rein, 1988), the large number of specimens, and the well known sedimentological (Aigner, 1985; Aigner and Bachmann, 1992; Aigner et al., 1999) as well as stratigraphic framework, specimens of these genera are excellent subjects for study. Their morphological fluctuations and dependence on palaeoenvironment, their mode of life (also discussed in Wang and Westermann, 1993; Rein, 1996, 1999a, 2000a; Klug et al., 2004), and immigrations into the German Muschelkalk Basin (Urlichs and Mundlos, 1985; Urlichs, 1999) are the focus of the following account. The only limiting factor of this study was the often moderate to poor preservation of the specimens, which made it difficult to find a statistically relevant number of sufficiently well preserved specimens. Immigration of ammonoids into the German Muschelkalk Basin has been documented for various genera, which are also known from the wider Tethyan realm (Urlichs and Mundlos, 1985; Urlichs, 1999): Acrochordiceras, Arcestes, Balatonites, Beneckeia, Bulogites, Discoptychites, Flexoptychites, Judicarites, Noetlingites, Paraceratites, Parapinacoceras, Protrachyceras (Nevadites according to Urlichs, 1999), and Trachyceras (Assmann, 1937; Balatonites according to Urlichs, 1999). A few nautiloid genera also immigrated into this epicontinental basin: Michelinoceras, Germanonautilus, Paranautilus, and Pleuronautilus. Some of the listed genera, such as Germanonautilus , have intruded the German

Muschelkalk Basin repeatedly. This genus is well known from both the Lower as well as the Upper Muschelkalk but there are no records from the Middle Muschelkalk; this can be explained by the hypersaline conditions during the timespan of deposition of the evaporites of the Middle Muschelkalk. In this study, we present the results of a stratophenetic analysis and the morphological fluctuations of the ammonoids of the Upper Muschelkalk (1), we discuss the relationships between changes in their morphology and changes in their mode of life (2), we review potential extinctions of ammonoid taxa of the Germanic Basin (3), and we report subsequent renewed immigrations of ammonoids (4).

2. Material Because of deformation or insufficient completeness of many specimens, approximately 500 specimens were selected out of several thousand. Only 274 of these specimens yielded satisfying data sets for the stratophenetic analysis. The measurements used in this study were taken from specimens that are deposited in the following institutions: The majority is stored in the Staatliches Museum fu¨r Naturkunde in Stuttgart (SMNS). A large number of specimens was made accessible at the Muschelkalkmuseum Hagdorn in Ingelfingen (MHI). Additional specimens were studied in the Museum fu¨r Naturkunde der Humboldt-Universit7t zu Berlin and the Geologisch-Pal7ontologisches Institut und Museum der Universit7t Tu¨bingen (GPIT). All measurements and computed values are listed in Appendix A. All specimens are internal moulds. Some phragmocones are incompletely filled; some contain micritic sediment, whereas others are filled with sparry cements.

3. Methods From 257 specimens, conch parameters and dimensions were measured (see Fig. 2). The measurements are listed in Appendix A. In order to obtain an idea of the morphological trends and pathways, univariate and multivariate statistical methods were applied. The simplest way to document trends is to plot morphological parameters trough time. Therefore,

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we computed the dispersal of different morphological parameters (diameter, whorl height/diameter ratio, umbilical width/diameter ratio, whorl expansion rate, whorl width/diameter ratio, and imprint zone rate; compare Korn and Klug, 2002) for each stratigraphic unit (Fig. 3). From the biometric data set (Appendix A), the following morphological characters were chosen for a multivariate statistical analysis: the whorl expansion rate (WER), whorl width index (ww/dm), whorl height index (wh/dm), umbilical width index (uw/dm), imprint zone rate (IZR), and features of the sculpture (sbc and sph, n, and sa; definition of characters see below). For a better understanding of the occupied morphospace, a canonical discriminant function analysis was carried out for each stratigraphic interval and for all measured specimens. In the first step, the values of the characters were standardised (z-standardisation). The discriminant factors for each specimen were extracted. F-values and Wilk’s Lambda were used to confirm the significance of the factors. Additionally, pooled within-group correlations between discriminating variables and canonical discriminant functions were calculated (see Schatz, 2001). The structure matrix and the eigenvalues are given in Appendix C. Three different data sets were used: (1) All data were used, including incomplete data sets. Missing values were interpolated. (2) Only the complete data sets were applied. The incomplete ones were erased. (3) Only the largest and best preserved specimen of each species was used. By applying this method, we made sure to include only adult specimens. This procedure bears the advantage that all specimens represent approximately the same ontogenetic stage. Additionally, juvenile and older preadult specimens of different taxa are morphologically less distinct.

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sediments of the Muschelkalk are of Anisian to Early Ladinian age (Aegean/Bithynian to Longobardian substage, 244 to 231 Ma according to Menning and the German Stratigraphic Commission, 2002). The Lower Muschelkalk belongs to the late Aegean/early Bithynian to early Illyrian substages, whereas the carbonates of the Upper Muschelkalk were correlated by Bachmann et al. (1999) with the late Illyrian to early Longobardian substages. For the canonical discriminant function analysis and the stratophenetic analyses of distinctive morphometric parameters, the Upper Muschelkalk was subdivided into 16 stratigraphic intervals (compare Urlichs, 1993) which are defined by the first occurrences (FO) of distinct ceratite species. Consequently, these units largely correlate with the currently accepted ceratite zonation in Germany. This subdivision is based on the range chart published by Urlichs (1993: 154). The subdivision (Fig. 1) is as follows (with the taxa names as given by Urlichs, 1993): 1—FO of Paraceratites (Progonoceratites) atavus atavus (Philippi, 1901); 2—FO of Pa. (Pro.) flexuosus flexuosus (Philippi, 1901); 3—FO of Ceratites (Doloceratites) pulcher Riedel, 1916; 4— FO of C. (Do.) robustus robustus Riedel, 1916; 5— FO of C. (Opheoceratites) raricostatus Riedel, 1916; 6—FO of C. (O.) compressus compressus Philippi, 1901; 7—FO of C. (O.) evolutus evolutus Philippi, 1901; 8—FO of C. (Acanthoceratites) spinosus spinosus Philippi, 1901; 9—FO of C. (A.) postspinosus Riedel, 1916; 10—FO of C. (Discoceratites) enodis (Quenstedt, 1849); 11—FO of C. (? A.) sublaevigatus Wenger, 1957; 12—FO of C. (C.) praenodosus Wenger, 1957; 13—FO of C. (C.) nodosus Schlotheim, 1820; 14—FO of C. (Di.) weyeri Urlichs and Mundlos, 1987; 15—FO of C. (Di.) dorsoplanus Philippi, 1901; 16—FO of C. (Di.) meissnerianus Penndorf, 1951.

5. Conch parameters 4. Stratigraphy The main focus of this study concerned the cephalopods from the Upper Muschelkalk. Consequently, the stratigraphy of the Lower Muschelkalk will not be discussed in detail here (Fig. 1). The

It is certainly the merit of authors such as Sandberger and Sandberger (1850–56) and later Trueman (1941), Raup (1966, 1967), as well as Saunders and Shapiro (1986) that they recognised the importance of the geometry of ammonoid coiling for systematics and ecology. Chamberlain (1976,

10

immigrants

ammonoids - bold italics, nautilids - italics brachiopods - underlined, crinoids - grey

Michelinoceras campanile C. enodis, C. laevigatus, Germanonautilus suevicus Coenothyris cycloides, Holocrinus doreckae ‘Protrachyceras recubariense’, ? Parasturia cf. emmerichi C. evolutus, Flexoptychites angustoumbilicatus, Punctospirella fragilis, Encrinus greppini P. atavus, G. bidorsatus Tetractinella trigonella En. liliiformis, Chelocrinus schlotheimi Arcestes sp.

Middle

Obere Sulfatschichten Salinar Fm. Steinsalzschichten Unt. Sulfatsch. Unt. Dolomit-Fm.

Lower

Pelson

Geislingen-Fm.

En. liliiformis, Entrochus silesiacus, Hol. dubius P. trinodosus, ‘C.’ antecedens Discoptychites dux, Judicarites stautei

Terebratelschichten Wellenkalk-Fm.

buchi-Mergel Oolithbank

Bithyn

Mosbach-Fm. 244

Geislingen-Bank Schaumkalkbank orbicularis Mergel

Wellendolomit Grenzkalkbank

Bulogites Balatonites spp., Beneckeia buchi Acrochordiceras damesii, Hol. dubius, En. ? radiatus Noetlingites strombecki, G. dolomiticus, G.salinarius, En. brahli, Moenocrinus deeckei, Dadocrinus sp. Ben. tenuis, Hol. beyrichi, Hol. wagneri, Dad. sp.

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immigrations

eustatic sea-level 3rd order cycles (Haq et al. 1988)

Upper

Illyr

Muschelkalk

Anisian

?

Unterer HauptMuschelkalk Obere Dolomit-Fm.

?

?

Oberer HauptMuschelkalk

curve after (Aigner et al. 1999) deeper marine shallow marine restricted marine coastal plain

?

Grenzbonebed 16 semipartitus Obere Terebratelbank 15 dorsoplanus Hauptterebratelbank nodosus 14 13 praenodosus sublaevigatus 12 enodis cycloides-Bank 11 Holocrinus-Bank 10 postspinosus spinosus 9 evolutus reticulata-Bank 8 ‘Spiriferina’-Bank 567 compressus robustus 4 pulcher Trochitenschichten 3 2 atavus Tetractinella-Bank 1 Hornsteinbank

large scale baselevel cycles (Aigner et al. 1999)

232

ceratite Zones (Urlichs 1993)

Fassan

?

regional units & marker beds

stratigraphic unit

Ladinian

Langobard

age in Ma (Hagdorn & Simon 2002)

substage

stage ?

?

lithostratigraphy

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

11

samax ah wh

OA

uw dm BCL ww = samin

Fig. 2. Conch parameters and dimensions. ah, apertural height; BCL, body-chamber length; dm, diameter; samin, whorl width between nodes, spines or ribs (=ww), samax, whorl width including ornament; uw, umbilical width; wh, whorl height; ww, whorl width. Modified after Klug et al. (2004).

1981), Jacobs (1992), and Jacobs and Chamberlain (1996) contributed important details on energy consumption, drag, the connection between conch size and flow resistance. 5.1. Whorl expansion rate In past decades, the whorl expansion rate proved to be a valuable proxy for the body chamber length and thus to some extent for the orientation of the aperture during life of the animal (Trueman, 1941; Raup, 1966, 1967; Saunders and Shapiro, 1986; Korn, 2000; Korn and Klug, 2001, 2002, 2003; Klug, 2001; Klug and Korn, 2004; Klug et al., 2004; Fig. 2). The orientation during life does not differ significantly from the post mortem orientation of a still gas-filled phragmocone (Reyment, 1980). Since the body chamber length correlates with the conch orientation, it also represents a proxy for the mode of life, swimming velocity, and manoeuvrability. Additionally, the whorl expansion rate reflects the overall geometry of the body chamber, which is of cardinal importance for the arrangement of musculature and other organs of the animal: Longidome forms probably had more elongate inner organs and the capability to retract arms and other temporarily external soft parts certainly differed from that of brevidome forms.

The equation to compute the whorl expansion rate, as suggested by Raup and Michelson (1965), Raup (1966, 1967), as well as Raup and Chamberlain (1967) uses measurements referring to the coiling axis (radii) of the ammonoid, which more or less crosses the protoconch perpendicular to the plane of symmetry [W=(d/e)2; Raup and Michelson, 1965; Saunders and Shapiro, 1986]. Since the internal whorls of many ammonoids had thin shells, they are not always preserved. This is especially true for the ammonoids from the German Muschelkalk Basin but also for many Palaeozoic forms. An estimation of the centre of coiling is thus difficult, and any misinterpretation causes a deviation from the correct value. This was the reason why Korn (2000) introduced an alternative algorithm for the whorl expansion rate, using diameters instead of radii [WER=(dm 1 / dm2)2=(dm/dm ah)2; Fig. 2]. This algorithm can be applied to any whorl fragment exceeding half a whorl, and yields almost identical values to Raup’s equations. Comparable expansion rates can be computed for the whorl width, the whorl height, and the whorl cross sectional area (Korn and Klug, 2001). 5.2. Whorl width index Similar to the WER, the whorl width index (ww/ dm) plays an important role for the ecology of

Fig. 1. Stratigraphic table of the German Muschelkalk (Anisian, Ladinian) with the large scale base-level cycles (modified after Aigner et al., 1999 and Menning and Deutsche Stratigraphische Kommission, 2002) and eustatic cycles (modified after Haq et al., 1988). Note that the curve from Haq et al. (1988) was correlated for each stage, i.e. there might be mistakes because of differing definitions of stage boundaries. Macroinvertebrate immigrants into the German Muschelkalk are indicated on the right (data from Urlichs and Mundlos, 1985; Hagdorn, 1985).

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ammonoids, especially with respect to drag and, consequently, swimming velocity (Chamberlain, 1976, 1981, 1987; Jacobs, 1992; Jacobs and Chamberlain, 1996). In addition to the WER and the umbilical width, the index defines the geometry of the body chamber and the arrangement of soft tissues, such as the length and diameter of muscles. In contrast to the diameters that are needed to compute the WER values, the whorl width is often impossible to be measured precisely, because the body chambers of many ammonoids from the Germanic Basin are deformed by compaction, corrosion, erosion, or were incompletely filled with sediment after deposition. Precise and correct whorl width values are therefore rare and can seem to be errors in the analysis. The mechanisms listed above are only responsible for a reduction of the conch thickness. Therefore, high values probably are closer to the true values than lower; exceedingly high values might actually be the best. This has to be kept in mind when reading the results of the analyses (Figs. 2 and 3). 5.3. Umbilical width index According to Raup and Michelson (1965), Raup (1966, 1967), and Raup and Chamberlain (1967), the umbilical width index (uw/dm) also correlates to some extent with the body chamber length, though significantly less so than with the WER. In contrast to Raup and his co-authors, we did not measure radii with reference to the coiling axis but the complete diameter. Among the forms of the Germanic Muschelkalk Basin, this parameter can be determined more reliably than whorl width because the umbilicus is often fairly well preserved.

the forms that were extracted from the Upper Muschelkalk, this diagenetic effect can be ignored. The facies is more or less calcareous throughout the entire Upper Muschelkalk, and most of these specimens are more or less adult (some display adult modifications). Therefore, the maximum size values for each stratigraphic interval should cause only a minor error. Conchs of presumably mature ceratites usually display an increased umbilical width, a smoother sculpture near the adult aperture, pronounced crowding of the terminal septa, narrowly spaced lirae near the adult aperture, and, rarely, they display constrictions that probably represent shell thickenings adjacent to the adult aperture. 5.5. Body chamber length The preservation of ammonoids from the Upper Muschelkalk often deteriorates towards their aperture; adult apertures are rarely well preserved (Philippi, 1901; Sun, 1928; Wenger, 1957). Nevertheless, in most specimens that are preserved with the body chamber, not much of it is missing. This fact became clear in an investigation of specimens exhibiting welldefined adult apertures, and it could also be tested in specimens where the black layer is preserved (a structure that is well-known from Recent Nautilus, where the dorsal mantle is attached to the shell; compare Klug et al., 2004). Most importantly, good estimates of the body chamber length can be obtained by using the correlation of conch geometry (especially the WER) with the measured body chamber length of complete conchs. 5.6. Sculpture

5.4. Maximum conch size Among ammonoids from the Germanic Basin, the maximum diameter changes significantly through time. In some cases, however, this parameter is biased by preservation of the material. Some localities, for example, yielded numerous specimens of the genus Beneckeia (early Anisian, Lower Muschelkalk) preserved in ferric oxides and sulphides. Usually, these do not exceed 15 mm in diameter. At numerous other localities, internal moulds of Beneckeia consisting of limestone were collected, which attain 10 cm. Among

The shell surface of ammonoids from the Germanic Basin varies from smooth (enodis/laevigatus group) to coarsely ribbed (nodosus group) to spinose (spinosus group). Due to the observation that the sculpture usually becomes weaker or even vanishes towards the (adult) aperture, the sculpture was measured at the posterior end of the body chamber. Additionally, any deterioration in preservation near the aperture is excluded by using this method. For this parameter, the whorl width occurring at (samax) and between the sculptural elements (samin) was

unit 11 unit 12

unit 13

unit 14 unit 15

unit 16

nodosus

weyeri dorsoplanus

semipartitus

unit 9 unit 10

unit 7 unit 8

unit 6

unit 5

unit 3 unit 4

sublaevigatus praenodosus

0.4 0.3

*

0.2 0.1

*

maximal value 75 % median value 25 % minimal value outlier value extreme value

*

* *

100

* * 10 3

0.4

whorl expansion rate

0.3

0.2

0.1

*

postspinosus enodis

semipartitus

weyeri dorsoplanus

nodosus

sublaevigatus praenodosus

postspinosus enodis

evolutus spinosus

raricostatus

compressus

pulcher robustus

0

evolutus spinosus

0.1

compressus

0.2

immigration 3

0.3 pulcher robustus immigration 2 raricostatus

immigration 3

immigration 2

immigration 1

atavus

0.4

flexuosus

*

0.3

flexuosus

*

immigration 1

*

0.4

0

1.5

0.5

imprint zone rate

0.5

0.1

2

1 0.6

0 0.6

0.2

*

2.5

atavus

umbilical width/diameter ratio

unit 2

0.5

0

whorl width/diameter ratio

13

1000

diameter in mm

whorl height/diameter ratio

0.6

unit 1

unit 16

unit 14 unit 15

unit 13

unit 11 unit 12

unit 9 unit 10

unit 7 unit 8

unit 6

unit 5

unit 3 unit 4

unit 2

unit 1

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Fig. 3. Fluctuations in the values of six major conch parameters of ammonoids during the deposition of the Upper Muschelkalk. Median values, the medium two quartiles, the maximum and the minimum value, outlier values as well as extreme values are indicated for each of the 16 stratigraphic intervals. The bold bars indicate the approximate timing of ammonoid immigration into the Germanic Basin.

measured and the numbers divided (sa=samin/samax). As mentioned regarding whorl width, the accuracy of these measurements can suffer from various tapho-

nomic and diagenetic alterations of the fossils. Only a few specimens yielded good values of this parameter.

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Additionally, the number of nodes or spines was quantified. Among the species from the Upper Muschelkalk, such elements are present in two positions, on the midflank and in the ventrolateral position. These two kinds of sculptural elements were counted (lateral: n l; ventrolateral: n vl) and divided (n=n l/n vl). Finally, the quality of the sculpture was grouped in six categories, smooth, smoothly dichotomous, strong dichotomous nodes, weak ribs, strong ribs, and ribs with prominent ventrolateral spines (Fig. 4). This list implies an increase in strength of the sculpture and was coded 0 to 5 in order to obtain an approximate quantification of the sculpture. Intermediate numbers represent intermediate states, e.g. 2.5. The quality of the sculpture was identified separately for the body chamber (sbc) and the phragmocone (s ph), because in many species, sculpture varies significantly within the terminal whorl.

5.7. Orientation of the aperture The orientation of the aperture can be identified mathematically with the algorithms of Raup and others (Raup and Michelson, 1965; Raup, 1966, 1967; Raup and Chamberlain, 1967; Saunders and Shapiro, 1986; Klug, 2001; Fig. 2). In contrast to many Palaeozoic forms, the whorl expansion rate and thus the body chamber length as well as the orientation of the aperture did not undergo extensive changes throughout ontogeny and phylogeny. Like many other Mesozoic ammonoids, most ceratites from the Germanic Muschelkalk Basin had body chamber lengths of approximately half a whorl. According to the authors listed above, the orientation of the aperture (OA) can be measured in degrees from the vertical direction. Most of the ammonoids from the Germanic Basin are oriented at 608 to 908 during life, so that the aperture was approximately horizontal during most of their lifetime.

Fig. 4. Ten characteristic species from the Upper Muschelkalk, arranged according to their stratigraphic appearance (left: the oldest species; right: the youngest species). The specimens are illustrated to scale (0.25) according to the median conch size of the stratigraphic unit listed below. The drawings were taken from Philippi (1901). The taxa are (from left to right; with the quality of the ornamentation in brackets): Paraceratites atavus atavus (smoothly dichotomous), Ceratites robustus terminus (strong dichotomous nodes), C. compressus compressus (moderately strong ribs), C. evolutus evolutus (strong ribs), C. spinosus spinosus (ribs with prominent ventrolateral spines), C. enodis (smooth), C. nodosus nodosus (strong ribs), Discoceratites weyeri (weak ribs), D. dorsoplanus (smooth), D. semipartitus (smooth). Note the changes in size and ornament, as well as the supposed immigrations (arrows).

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Rein (1996, 1999a, 2000a) obtained similar results for the orientation of the aperture but he argued that this would have caused a permanent downward movement of these ammonoids, because the aperture was higher than the centre of gravity and the excurrent of the hyponome acted on the centre of gravity. This assumption, however, completely neglects the flexibility of the hyponome (Packard et al., 1980) as well as the role of drag at a high swimming velocity (Klug and Korn, 2004). Additionally, Rein (1996, 1999a, 2000a) denied the ability of the ammonoids from the Upper Muschelkalk to maintain neutral buoyancy. His hypothesis was based on his own calculations and those by Ebel (1990). However, their conclusions are questionable for the following reasons. (1) Ammonoids from the Germanic Basin are never preserved with shells replaced by calcite or silica. Consequently, the shell thickness (which is obligatory for buoyancy calculations) can only be guessed and not precisely measured. This does not, of course, prove the ability of these Triassic ammonoids to achieve neutral buoyancy but it demonstrates the limitations with respect to the accuracy of buoyancy calculations. Jacobs and Chamberlain (1996: 185) commented, bit is surprising that the results of Ebel [. . .] are as close to neutral buoyancy as they are. These results could just as easily be interpreted as a confirmation of neutral buoyancy.Q (2) The WER values of Triassic ceratites show a universal correlation to the body chamber lengths of other Mesozoic ammonoids. If the function of the phragmocone was not buoyancy, it probably would not have undergone such changes during the phylogeny of these ammonoids or it would have been reduced to some extent. (3) Ceratites which carried syn vivo epibionts, such as the oyster Placunopsis, reacted to these epizoans by changing the growth of the body chamber while keeping up the growth of the septa (for a differing opinion see Rein, 1996). Consequently, the body chamber became shorter but wider (increased WER) in some of these specimens. For a fully epibenthic organism, there would have been no need to change the mode of growth in such a way as a reaction to dhitchhikersT such as Placunopsis (Meischner, 1968; Seilacher, 1960; Davis et al., 1999; Klug et al., 2004).

15

6. Morphological fluctuations 6.1. General The canonical discriminant function analysis revealed a strong discrimination along the first function (Appendix C). The function 1 represents 68.7% of overall variance. Structure features (sa, sbc and sph) especially correlate with this function. The parameters correlating with function 2 (18.2%) are associated to the parameters WER and ww/dm; these are parameters that are in a context to the orientation of the aperture during life (see above). The interpretations of the morphological fluctuations are based on scatter plots (Figs. 5 and 6) representing nearly 87% of the variance. For a general overview of the fluctuations in morphospace occupation, canonical discriminant function analyses were carried out for the 16 stratigraphic units. The graphs of the three data sets listed in the methods section are more or less similar in their overall pattern (Fig. 5). A more pronounced grouping of the data sets of the centroids from each stratigraphic unit along the second function is the main difference between the graph with all specimens and those with only the complete sets or only the largest specimens of each species. This arrangement in groups of the centroids of three to five stratigraphic levels is largely the same in the three graphs (Fig. 6): The centroids of levels 1 to 4, 10, 15 and 16 are grouped together (morphogroup A), the centroids of the units 5 to 7 with 11 and 14 are close to each other (morphogroup B), and the centroids of the units 8, 9, 12 and 13 are quite close (morphogroup C). In any case, the morphological change is most pronounced from the stratigraphic level 9 to 10. The morphological pathway, beginning at morphogroup A and ending in morphogroup D, describes two more or less congruent horizontal elliptical paths, crossing the morphogroups in the order A–B–C–A– B–C–B–A. This ordering in morphogroups is only helpful for the general pattern. For a more detailed analysis, the changes between the single levels 1 to 15 had to be studied. It is not surprising that the species of the stratigraphic units 1 to 4 are grouped together because they have a dichotomous sculpture which

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C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

morphological fluctuations of the ceratitids of the Germanic Basin (late Anisian to early Ladinian)

6 function 2

A

4

2

0

-2

-4

6 function 2

B

4

2

0

-2

-4 -6

-4

-2

0

2

function 1

4

Fig. 5. Results of the canonical discriminant function analyses with two of the three data sets: A. All specimens with interpolated missing values; B. only the largest and best preserved specimens. The three bold arrows mark the approximate direction of the evolutionary pathways in between the three immigrations and the respective extinctions. The species figured next to some of the centroids are 1: Paraceratites atavus atavus; 6: C. compressus compressus; 7: C. evolutus evolutus; 8: C. spinosus spinosus; 10: C. enodis; 13: C. nodosus nodosus; 16: D. semipartitus. They are approximately illustrated to scale (0.17) and were taken from Philippi (1901). The function 1 represents 68.7% of overall variance, whereas function 2 represents 18.2% of the variance.

function 2

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34 6

morphological fluctuations of the ceratitids of the Germanic Basin (late Anisian to early Ladinian) data: all specimens of all species with interpolated ‘missing values’ n = 274

4

2

1

extremevalues of groups 4, 6

A 13

2

12

4 3

6

9

4

15

0

C

8

16 10

6

14

7

11 5

-2

B

preadult specimens of groups 8, 9

-4 -6

-4

-2

0

2

4 function 1

Fig. 6. Results of the canonical discriminant function analyses with all specimens with interpolated missing values. The shaded areas indicate the morphogroups A, B, and C. Note the gap between A and B that is only bridged by one extreme value each of the stratigraphic units 4 and 6. The vertical oval indicates the position of mainly pre-adult specimens of the units 8 and 9, in which the characteristic sculpture is not fully developed. The specimens of unit 15 remarkably fall both in the morphogroups A and B; this can be explained by the presence of specimens which retained remains of the sculpture characteristic of morphogroup C. The three bold arrows mark the approximate direction of the evolutionary pathways in between the three immigrations and the respective extinctions.

increased in strength during phylogeny. Their ancestor–descendant relationships are rather clear (e.g. Urlichs and Mundlos, 1980; Urlichs, 1993, 1999). The taxa that were extracted from the stratigraphic intervals 5 to 7 were grouped in the subgenus Ceratites (Opheoceratites) by Schrammen (1928; see also Urlichs and Mundlos, 1987) because of their coarse ribs in the adult growth stages and the comparatively wide umbilicus. During phylogeny of this subgenus, the dichotomous phase ended earlier and coarse ribs were formed earlier in ontogeny. The close morphological relationships between the ammonoid taxa from the units 8 and 9 are also reflected by their grouping in the subgenus Ceratites (Acanthoceratites). They share the presence of long ventrolateral spines in the adult whorl.

17

The morphological step from unit 9 to 10 is clearly the most extreme. Unit 10 contains mainly two forms, Ceratites enodis and Ceratites laevigatus. Both have a smooth to weakly ornamented adult body chamber. The latter species occasionally has a few ribs near the anterior end of the phragmocone, similar to Ceratites sublaevigatus or Ceratites nodosus. This phenomenon can also be seen in various species of Discoceratites (D. alticella, D. meissnerianus, and D. weyeri). These sculptural elements are slightly stronger developed in the forms of unit 11 and much stronger in those of units 12 to 14. Nevertheless, the sculpture is certainly the strongest near the transition of the phragmocone to the body chamber of the adult specimens, and hence probably reflects a distinct feature of the terminal growth. Towards the aperture of the adult body chamber, the sculpture becomes more or less smooth again. In the units 15 and 16, only representatives of Discoceratites were found. Some species of this genus retained coarse ribs near the last septa, but in all species, the sculpture becomes smooth towards the adult aperture. All taxa of the latter two stratigraphic units have large conchs (dm up to 400 mm), they are discoconic, and have a narrow umbilicus. Only a few ammonoids (Alloceratites schmidi, Neoclypites peregrinus; Urlichs, 1999) were extracted from the Lettenkeuper (Late Longobardian, Late Ladinian). These two forms differ completely in their morphology from the members of Discoceratites and thus, they are probably not directly related to them. No ammonoid remains were discovered yet from the interval between the last occurrence of Discoceratites and the first of Alloceratites and Neoclypites. Supposably, this represents another transgression and an immigration from the Tethyan realm (Urlichs, 1999). The most important morphological trends among the ammonoids from the Upper Muschelkalk are: (1) the decrease of whorl height from unit 1 to unit 7 followed by an increase from unit 8 to unit 16; (2) a markedly expanding umbilical width until unit 5 and a subsequent decline, lasting until unit 16; (3) a general increase in size, with a decrease after unit 9; (4) a rise in the strength of the sculpture from unit 1 to 9 and again from unit 10 to 13, and finally a decrease until unit 16; (5) a clear disturbance in the morphological development at the transition from unit 9 to unit 10 in most parameters.

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

The whorl width index varies slightly from units 1 to 9 (median values of the whorl width index between 0.29 and 0.34; Fig. 3); the total range is between 0.19 and 0.53. From the stratigraphic unit 9 to unit 10, a striking decrease in the median ww/dm ratio from 0.35 to 0.27 can be observed. Except for the members of the subgenus Discoceratites from unit 16 and ceratites from the Lower Muschelkalk, these are the forms with the narrowest conchs. After unit 10, the whorl width index increased to a median value near 0.4 in the units 12 and 13 and is reduced to 0.25 in unit 16. Generally speaking, the ammonoids which reached the German Muschelkalk Basin in Anisian and Ladinian times had oxyconic (Beneckeia, Discoptychites, Flexoptychites, Noetlingites, Paraceratites, Parapinacoceras) or platyconic conch shapes (Acrochordiceras, Balatonites, bCeratitesb antecedens, Judicarites, Nevadites, Alloceratites; for references see Urlichs and Mundlos, 1985, or Urlichs, 1999). Truely globose or spheroconic forms (except for one specimen of Arcestes) either did not reach this basin or did not develop from one of the Tethyan immigrants within the basin; the whorl width index rarely exceeds 0.5. 6.4. Umbilical width index In contrast to the median whorl width index, the median umbilical width index displays a steady increase from stratigraphic unit 1 (0.17) to unit 5 (0.33). It stays high until unit 7, but subsequently,

From stratigraphic unit 1 to unit 16, the terminal diameter of the ceratites from the German Muschelkalk Basin increases almost constantly (Figs. 3, 5 and 7). In units 4 to 8, the median diameter reaches almost 100 mm. The ammonoids from unit 9, however, are clearly larger with a median diameter of 152 mm, and in the subsequent units 10 to 12, most ammonoids reach a diameter of approximately 100 mm. Most stratigraphically younger forms are significantly larger, with diameters of over 400 mm in unit 16. 6.6. Body chamber length According to Wenger (1957: 62), the body chambers of most ammonoids from the Upper Muschelkalk comprise two fifths to half a whorl (i.e. 1448 to 1808). According to measurements by Klug et al. (2004), the body chambers of these species are 1208 to 1908 long. The formation of short body chambers (b 1508) might have been caused by the encrustation by epizoans, usually by the oyster Placunopsis. Wenger (1957: 68)

300

100 90 80 70

250 200

60 50 40 30

150 100 50

diameter % of ceratitids with Placunopsis

0

20 10 0

% of ceratitids with Placunopsis

6.3. Whorl width index

6.5. Maximum conch size

unit1 unit2 unit3 unit4 unit5 unit6 unit7 unit8 unit9 unit10 unit11 unit12 unit13 unit14 unit15 unit16

In contrast to primary expectations, the fluctuations in the whorl expansion rate through time are rather low (Fig. 3); the majority of the ceratites have WER values between 1.7 and 2.3. The median value of this parameter is reduced from approximately 2.2 in unit 1 to 1.8 in unit 5, increased to 2.1 until unit 8 and 9 and shows a decrease towards unit 10 (1.9). It increased slightly until unit 16 (2.1). As discussed above, this implies only slight changes in body chamber length and also in the orientation of the aperture (see also below).

this ratio continuously decreased until unit 16, where it reaches the same median value as in unit 1: 0.17 (Fig. 3).

atavus flexuosus pulcher robustus raricostatus compressus evolutus spinosus postspinosus enodis sublaevigatus praenodosus nodosus weyeri dorsoplanus semipartitus

6.2. Whorl expansion rate

diameter in mm

18

Fig. 7. Median conch size (own data) and percentage of encrustation of the ceratitid conchs by the oyster Placunopsis ostracina (data from Wenger, 1957). The graphs show a good correlation of the conch size and the oyster encrustation.

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counted the numbers of ammonoid specimens with such encrustations. Among the species he studied, the percentage of specimens with Placunopsis varied between 0% and 82% with an average of 21% (Fig. 7). He noticed that the number of infested specimens increased more or less constantly from the oldest to the stratigraphically youngest taxa. One reason for this phenomenon might be the general conch size increase during this time interval. Wenger did not discuss whether these encrustations happened syn vivo or post mortem and consequently, he did not consider possible effects on the mode of growth of the body chamber and the buoyancy apparatus, respectively. 6.7. Orientation of the aperture Since the whorl expansion rate of the ammonoids from the Upper Muschelkalk does not vary extremely (1.7 to 2.3), the body chamber length was probably also rather uniform (mostly between 1508 and 1808). Consequently, the orientation varied between approximately 608 and 908 from the vertical. 6.8. Sculpture Ammonoids with oxyconic to thinly platyconic conchs with smooth or at least faint sculpture can be found in the Lower and Upper Muschelkalk. Among the older taxa, representatives of Beneckeia, Discoptychites and bCeratitesQ antecedens belong to this group. Among the taxa from the Upper Muschelkalk, there are three groups of ceratites which have such a morphology, i.e. the subspecies of Paraceratites atavus, the Ceratites enodis and Ceratites laevigatus bgroupQ, and most species of Discoceratites (Fig. 4). This is reflected in the values of the sa-ratio (in this case with values close to 1) and also of the n l/n vl ratio (values approximate 0.5). The conchs of the representatives of the subspecies of Acanthoceratites spinosus as well as of Ceratites nodosus including their closest relatives possess the strongest sculpture among the forms from the Upper Muschelkalk. The former have strong ventrolateral spines and the latter display coarse lateral ribs. In general, there is a tendency from weaker to stronger sculpture from the stratigraphic unit 1 to unit 9, and, after the sculpture was largely reduced in the conchs of the species of

19

unit 10, it increased again from unit 11 to unit 13. In subsequent units, the shell surface again became more or less smooth.

7. Physical conditions (sea-level) Changes in the sea-level within the Germanic Basin are documented in the Upper and Lower Muschelkalk in repeated changes of carbonate content, grain size, faunal composition, and other factors (see Aigner, 1985; Fig. 1). During Triassic times, the sedimentary environment changed from largely terrestrial or fluvial from the earliest Indusian to the earliest Anisian to shallow marine during most of the Anisian and Ladinian (with a phase of evaporite deposition in the late Anisian) to predominantly terrestrial and fluvial from the late Ladinian to the end of the Triassic. For the deposits of the Upper Muschelkalk (late Anisian and early Ladinian), one large scale base-level cycle was documented by Aigner et al. (1999). This event can be subdivided into two smaller cycles, with rather shallow marine conditions during the formation of the sediments of the stratigraphic units 1, 4, 15, and 16 and with deeper marine conditions during deposition of the carbonates of units 2 to 3 and 8 to 14 with a maximum in unit 10 (Fig. 1). Compared to the curve published by Aigner et al. (1999), the eustatic sea-level curve produced by Haq et al. (1988) differs quite considerably (Fig. 1). Only the major trends within the two curves roughly coincide.

8. Environment In previous decades, various workers dealt with the marine palaeoenvironment during Middle Triassic times in the Germanic Basin from both palaeontological and sedimentological points of view (e.g. Aigner, 1985; Hagdorn, 1985; Hagdorn and Simon, 1993; Ockert, 1993; Aigner et al., 1999; Beutler and Szulc, 1999; Fig. 1). In contrast to the evaporites of the Middle Muschelkalk, the carbonates of the Lower and the Upper Muschelkalk were largely deposited under euhaline conditions. Fluctuations in salinity probably caused the short-lasting appearances of various organisms in the Germanic Basin, such as

20

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the mostly stenohaline ammonoids (Urlichs and Mundlos, 1985) and crinoids (Hagdorn, 1985; Hagdorn and Gluchowski, 1993). These salinity changes might well have been the cause for the extinction of some ammonoid taxa on the one hand and enabled the widespread settlement of the conchostracan Palaeestheria (almost immediately after the extinction of the spinosus-group) on the other hand (Urlichs, 1999). Contemporaneous with the salinity changes during the Anisian and Ladinian, the water depth changed. This is reflected in changes of grain size and sedimentary structures such as wave ripples, cross-bedding, alignment of shells, hardgrounds, bonebeds, reworking, etc. Sediment consistency varied from soft to hard, but the abundance of endobenthic organisms documents the predominance of softgrounds. A palaeolatitude of approximately 308 North in combination with the presence of coral reefs of Pelsonian age in Upper Silesia (Beutler and Szulc, 1999) implicate a warm to temperate climate for the Germanic Basin. Powerful storms are recorded in numerous tempestitic beds (Aigner, 1985) and currents are documented by various pot and gutter casts as well as the alignment of shells. Urlichs and Mundlos (1985) as well as Urlichs (1999) explained the presence of some cephalopods in the Lower Muschelkalk in the western part of the basin and that of the orthocone Michelinoceras in the Upper Muschelkalk by these currents, which carried empty conchs into the Germanic Basin. The effect of currents on the immigration of organisms into the Germanic Basin has not yet been studied in detail. Since ammonoids, including those of the Germanic Basin, were not arduous and fast swimmers, it appears likely that, at least to some extent, favourable currents supported the immigrations of ammonoids.

9. Migrations and palaeobiogeography During the early Anisian, marine conditions expanded from the easternmost Germanic Basin towards the west. Various authors have documented the subsequent immigration of organisms into this peri-Tethyal epicontinental basin (Assmann, 1944; Kozur, 1974; Senkowiczowa, 1975; Hagdorn, 1985; Hagdorn and Gluchowski, 1993; Kedzierski and Szulc, 1996; Beutler and Szulc, 1999; Urlichs, 1999; Fig. 8). A number of ammonoid taxa arrived particularly in the eastern part of the Germanic Basin during Bithynian to early Illyrian times (Acrochordiceras, Balatonites, Beneckeia, Bulogites, Discoptychites, Judicarites, Noetlingites, Paraceratites, Parapinacoceras, Protrachyceras; Noetling, 1880; Wurm, 1914; Rassmuss, 1915; Assmann, 1926, 1937; von Pia, 1930; Schmidt, 1934; Trammer, 1972; Kelber, 1977; Urlichs and Mundlos, 1985; Dzik, 1990; Hohenegger and Tatzreiter, 1992; Kaim and Niedzwiedzki, 1999; Urlichs, 1999; Niedzwiedzki et al., 2001). According to their occurrences, most of these genera probably entered this region via the Silesian–Moravian and the East Carpathian Gate. For bCeratitesb antecedens, however, a western entry was suggested by Schmidt (1934) because of its moderate abundance in the southwestern part of the Germanic Basin. To our knowledge, no ammonoid or brachiopod remains were discovered yet in the Middle Muschelkalk of the Germanic Basin. Finds of crinoids are restricted to the easternmost part of the Muschelkalk Basin near the Silesian–Moravian Gate (Hagdorn, 1985). The majority of the basin was too hypersaline for most marine macro-organisms to survive. In late Illyrian to Longobardian times, the majority of cephalopods had probably intruded the Germanic Basin through the Burgundian Gate (Urlichs and

Fig. 8. Eight palaeogeographic maps of central Europe with the outlines of the Germanic Basin during the Anisian and Ladinian with its connections to the Tethyan realm. The approximate occurrences of the ammonoids Paraceratites atavus atavus, Ceratites spinosus s.l., C. enodis, and C. laevigatus, of the brachiopods Tetractinella trigonella and Coenothyris cycloides as well as of the crinoids Encrinus liliiformis, Holocrinus doreckae are indicated in the respective maps (map modified after Hagdorn, 1985; distribution of the eight taxa modified after Hagdorn, 1985; Urlichs and Mundlos, 1985; Hagdorn and Simon, 1993; Urlichs, 1997, 1999; Niedzwiedzki et al., 2001 and own data). The spatial distribution of these taxa within the Germanic Basin suggests that they immigrated via the Burgundian gate (arrow 3). The maps in A to D display the distribution of the respective species in the atavus zone, those in E, G, and H show the distribution in the enodis/laevigatus zone and the map in F pictures the distribution in the spinosus zone; note the wider distribution of C. spinosus s.l. (F) compared to that of C. enodis and C. laevigatus (G). The question marks at the southern edge of the distribution of Ceratites spinosus s.l., C. enodis, and C. laevigatus indicate the presence of similar or identical forms in the Muschelkalk of the Toulon area in southeastern France (compare Urlichs, 1997, 1999).

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

A

North Sea e

e

2

2 Danube

area of nondeposition area Tethys of deposition

North Sea e

3

D

Germanonautilus bidorsatus

Elb

North Sea e

1

e

1

e

s ma

siv

Rhine

n

cia

eli

Tetractinella trigonella

Elb

siv

Rhine

0°

Muschelkalk Danube

front of alpidic deformation

d Vin

Tethys

Rhine

Rhine

n

cia

eli

s ma

Paraceratites atavus atavus

Elb

siv

d Vin

C

North Sea

1

e

3

B

Encrinus liliiformis

Elb

21

2

n

cia

eli

d Vin

Danube

s ma

2 Danube

Tethys 3

E

Tethys 3

front of alpidic deformation

North Sea

F

Holocrinus doreckae

Elb

e

North Sea e

1

e

siv

2

n

cia

eli

d Vin

Danube

s ma

Tethys 3 North Sea

3

H

Ceratites enodis/ C. laevigatus

Elb

e

2

Elb

e

3

Coenothyris cycloides

1

Danube

Tethys front of alpidic deformation

front of alpidic deformation

North Sea

Rhine

n

cia

eli

d Vin

?

1

e

siv

s ma

2 Danube

Tethys

front of alpidic deformation

Rhine

G

1

e

s ma

Rhine

Rhine

n

cia

eli

Ceratites spinosus s.l.

Elb

siv

d Vin

front of alpidic deformation

3

migration of faunal elements 1 East Carpathian gate 2 (early Anisian) 2 Silesian Moravian Danubegate (early Anisian) 3 Burgundian gate (late Anisian to early Ladinian)

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C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Mundlos, 1980, 1985; Urlichs, 1999). According to Senkowiczowa and Szyperko-Sliwczynska (1975), the Silesian–Moravian Gate was then closed and the East Carpathian Gate reopened but to a lesser extent than before. Remarkably, several Germanic species are also known from the Tethyan realm [e.g. Ceratites robustus robustus from Romania: von Pia, 1930; Wenger, 1957; C. enodis from southeastern France: Urlichs, 1997, 1999; C. (Acanthoceratites) sp., C. (Opheoceratites) sp., ’C. muensteri’, C. cf. nodosus from Sardinia: Ma´rquez-Aliaga et al., 2000: 104; Discoceratites cf. dorsoplanus, D. semipartitus from Bulgaria: Entcheva, 1972; further ammonoid associations with Germanic affinities have been reported from Israel, Spain, and even Nevada: Schmidt, 1935; Virgili, 1958]. Silberling (1959: 13) strikingly reported some discoveries of dGermanicT cephalopod forms such as Paraceratites cf. flexuosus, Discoceratites cf. dorsoplanus, D. cf. semipartitus and Germanonautilus cf. bidorsatus from the Grantsville Formation of the Shoshone District (Nevada). Apparently, these fossils are poorly preserved and he did not figure them. Thus, this record is somewhat dubious. In addition to Paraceratites, which gave rise to several taxa of the Upper Muschelkalk in the Germanic Basin, Arcestes, Flexoptychites, Nevadites, and Protrachyceras were discovered in the stratigraphic unit 1 (Urlichs and Mundlos, 1980, 1985; Urlichs and Kurzweil, 1997). Among other marine organisms which have entered the Germanic Basin from the Tethys Sea, the brachiopods and the crinoids are worth mentioning. In the Upper Muschelkalk of Germany, a number of stratigraphically important horizons (bO¨ kostratigraphische Leitb7nkeQ, Hagdorn and Simon, 1993) were named after their brachiopod content: bTetractinella-BankQ [Tetractinella trigonella (Schlotheim, 1820)], bSpiriferina-BankQ [Punctospirella fragilis (Schlotheim, 1813)], bcycloides-BankQ [Coenothyris cycloides (Zenker, 1836)], QBank der kleinen TerebratelnQ, QHauptterebratelbankQ and QObere TerebratelbankQ [the latter three with Coenothyris vulgaris (Schlotheim, 1820)]. It is likely, that the first three of these horizons reflect immigrations of brachiopods into the Germanic Basin and roughly correspond to comparable immigrations of certain ammonoids: Shortly after the return of euhaline conditions during the late Illyrian, Tetractinella

trigonella and Encrinus liliiformis Lamarck, 1801 spread across almost the entire basin, followed by the ammonoid Paraceratites atavus atavus soon thereafter; the morphological change among the ceratites from stratigraphic units 4 to 5 closely precedes the reappearance of Punctospirella fragilis, which inhabited the Germanic Basin already during the deposition of the Lower Muschelkalk, and of the crinoid Encrinus greppini De Loriol, 1877 (Hagdorn, 1985: 249); the ammonoids Ceratites enodis and Ceratites laevigatus as well as the brachiopod Coenothyris cycloides and the crinoid Holocrinus doreckae Hagdorn, 1983 appeared almost synchronously in most places within the Germanic Basin (Hagdorn, 1985; see also Rein, 2000b). Particularly the last case is, in our opinion, a good example for an immigration into this epicontinental basin, probably enabled by a transgression (compare Aigner et al., 1999). Coenothyris vulgaris lived in the Germanic Basin throughout most of the Anisian and Ladinian intervals and is thus of no use for this discussion. Presuming the correctness of the hypothesis of repeated extinction and immigration of ceratites in the Germanic Basin, the question for the origin of the ancestors of the immigrants remains to be answered. Even for the first immigration, this problem is not trivial. According to Urlichs and Mundlos (1985) and Urlichs (1999), Paraceratites atavus atavus migrated into the basin via the Burgundian gate. Urlichs and Mundlos (1980) suggested Paraceratites abichi (Mojsisovics 1882) (dSchreyerites abichiT in Tatzreiter and Balini, 1993) as its ancestor. For the later immigrations, no suggestions were previously made. Most early Ladinian ammonoids from the Tethys possess a distinct keel, whereas such a keel is absent in the forms from the Germanic Basin. Some records from the Ladinian deposits of the Sephardic Province, however, were assigned to species which were described using material from Germany (Stefanov, 1932; Virgili, 1958; Budurov et al., 1993; Goy, 1995; Ma´rquez-Aliaga et al., 2000; Posenato, 2002; Urlichs and Posenato, 2002). This extends the geographical range of some Germanic ceratite taxa of Ladinian age far outside of the German Muschelkalk Basin to Bulgaria, Italy, and Spain. Nevertheless, the precise process of regionally disappearing taxa and renewed migration to this basin (or maybe just the majority of the basin) remains

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unclear. In contrast to the above model, a rootstock of ceratites resembling members of the genus Paraceratites might have survived in some relic areas of the Germanic Basin or as well in the northern Tethys.

10. Ecology 10.1. Buoyancy Like most fossil ectocochleate cephalopods and like all Recent ectocochleate cephalopods, most ammonoids were most likely capable to achieve neutral buoyancy (see review in Jacobs and Chamberlain, 1996). This was doubted by some authors (Ebel, 1990; Rein, 1996, 1999a, 2000a). As discussed by Jacobs and Chamberlain (1996) and in the methods chapter, these doubts appear to be not very reasonable for the following reasons: (1) Buoyancy calculations come either close to neutral buoyancy or support the hypothesis for neutrally buoyant ammonoids (Jacobs and Chamberlain, 1996). (2) The ammonoid phragmocone underwent an intricate evolution which is hardly explainable if it was not functional at all. (3) Ammonoids reacted on epizoans in various ways including the reduction of body chamber length or asymmetric growth (Klug et al., 2004). This can hardly be explained by a benthic mode of life. (4) Soft-tissue attachment structures strongly resemble those of Recent nautilids (Klug, 2004; Klug and Lehmkuhl, 2004; Klug et al., 2004; unpublished material) and hence, a similar, at least partially nektonic mode of life appears reasonable. Therefore, ceratites were probably also able to achieve neutral buoyancy. 10.2. Shell shape and swimming velocity Ceratites mostly have brevidomic platyconic to oxyconic conchs. According to Westermann (1996) this implies a more or less nektonic mode of life. The strength of the sculpture varies considerably from species with completely smooth conchs to species with long ventrolateral spines or strong ribs. Strong ornament certainly increased drag and thus, the maximum swimming velocities varied probably considerably between the various species of Ceratites, Discoceratites and Paraceratites. According to the review on drag and swimming by Jacobs and

23

Chamberlain (1996), the large oxyconic representatives of Discoceratites with smooth conchs were good swimmers at higher velocities and also at accelerating. The less compressed involute to evolute platyconic species of Ceratites and Paraceratites were stronger in continuous swimming at low velocities but accelerated slower. For discoconic, platyconic and oxyconic brevidomes, such as the ceratites from the Germanic Basin, Westermann (1996) suggested a very good swimming potential for backward motions combined with a moderate to good steerage. 10.3. Shell orientation and manoeuvrability The approximate body chamber lengths of the brevidomic ceratites are fairly well known (e.g. Philippi, 1901; Sun, 1928; Wenger, 1957; Klug, 2004). In most specimens of Ceratites, Discoceratites and Paraceratites, the body chamber measures between 1448 and 1908, with the majority near 1808. Depending on shell thickness, on shell geometry and also, to some minor extent, on adult modifications (Klug, 2004), the orientation of the aperture of most ceratites was 608 to 908. In most ceratite species, the aperture was close to a horizontal position. Consequently, the position of the opening of the hyponome was close to the maximum horizontal distance from the centre of gravity. This implies that steering was very good in ceratites. 10.4. Epizoans True epizoans can only rarely be identified in the fossil record (Davis et al., 1999). In fact, many cases proved to be dbenthic islandsT (Seilacher, 1981) when examined closer. Considering ceratites and the oyster Placunopsis, Placunopsis can be shown to have lived both as true epizoan and on dbenthic islandsT (e.g. Wenger, 1957; Rein, 1996, 1988, 1999a, 2000a,b; Klug and Lehmkuhl, 2004; Klug et al., 2004). In several specimens, Placunopsis was overgrown by succeeding whorls of the ceratite, proofing the epizoan nature of this oyster. Additionally, some ceratite specimens were embedded vertically with the body chamber at the bottom. Sometimes, these specimens are preserved with Placunopsis with conjoined valves. This is of course no proof but at least an indication for syn vivo encrustations. It is important to repeat at

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C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

this point that the body chambers of some ceratite specimens carrying Placunopsis valves actually have shorter body chambers or show asymmetric growth. This is also a strong argument for syn vivo encrustation. For most finds of this oyster on ceratites, however, it is admittedly difficult to impossible to find true evidence that Placunopsis has inhabited the living ceratite. For a list of further organisms that inhabited ceratite shells and references on that subject see Klug and Lehmkuhl (2004). 10.5. Sexual dimorphism Sexual dimorphism is a phenomenon which is well documented for many Jurassic and Cretaceous ammonoids (for a review see Davis et al., 1996; important papers: Makowski, 1962; Callomon, 1963; Westermann, 1964). For Triassic ammonoids, only a few reports are available (e.g. Dzik, 1990). The same is true for ammonoids from the Germanic Basin (Rein, 1999b). Interestingly, the latter author distinguished the dimorphic forms based on smooth versus sculptured internal whorls. This differs from the criteria usually applied for dimorphic pairs among the Ammonoidea (morphologic distinction, identical early ontogeny, identical stratigraphic ranges; see the articles by the authors listed in the first sentence of this paragraph). It appears still reasonable to assume sexual dimorphism for ceratites. In fact, the low number of species and the good stratigraphic control make the ceratites suitable study objects for sexual dimorphism. The only problem is the poor preservation, especially of the internal whorls, making the study of their early ontogeny hard to impossible. Yet, when regarding the ceratite range chart published by Urlichs (1993), however, most ceratite zones contain two to five species and subspecies. In many of these cases, one (or two) species displays strong ornament and the other one (or two) is smooth or less strongly ornamented. This might be an indication for sexual dimorphism in ceratites from the Germanic Basin requiring further examination. 10.6. Salinity During Anisian to Ladinian times, the salinity of the sea water in the Germanic Basin changed significantly. Limestones, dolomites, gypsum, anhydrites, and vari-

ous salts were deposited. In consequence, diversity of marine life varied extremely from no traces of fossils in the evaporites to more or less diverse faunas in the limestones. The limestones of the Lower and Upper Muschelkalk contain both stenohaline (echinoderms, articulate brachiopods) and euryhaline forms (various bivalves like e.g. Myophoria, Placunopsis). It is remarkable, however, that simultaneous with ammonoids, some stenohaline forms migrated into the Germanic Basin. This is an indication for the stenohaline preferences of ceratites, thus contradicting the suggestions by Wang and Westermann (1993) and Westermann (1996). Additional support of this hypothesis was given by Urlichs (1999), who wrote that a phase with hypersaline conditions and an absence of ammonoids in most of the Germanic Basin followed after the extinction of the spinose forms. Freshwater input is documented in the composition of arthropod faunas of this stratigraphic interval in the northern part of the Germanic Basin (Kozur, 1974; Urlichs, 1999). Hence, the ceratites were probably stenohaline like most other ammonoids.

11. Conclusions Our stratophenetic case-study on the fluctuations in morphometric parameters of cephalopods through time shows how regional biodiversity in a marine epicontinental basin can be affected by sea-level changes (compare Monnet et al., 2003). Discontinuities in the morphological change rather reflect smallscale extinctions and iterative immigrations than a purely endemic evolutionary process which was restricted to this basin. This hypothesis is supported by the repeated immigrations of other faunal elements (e.g. crinoids and brachiopods), which are approximately synchronous with the morphological discontinuities in the evolution of ammonoids that occurred through time. In some cases, these postulated immigrations coincide with sea-level highstands (Fig. 1). This implies a widened and deepened passage between the restricted basin and the ocean and possibly decreasing salinity and thus recovering conditions for marine life. In this case, a sea-level rise caused a regional diversity rise. It is well possible, however, that in between these discontinuities, new species of ceratites have evolved within the Germanic

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

basin. Especially the forms of the stratigraphic units 5 to 9 and 12 to 16 display rather gradual morphological transformations. The ecology of ceratites can be summarised as follows: (1) Ceratites normally were able to achieve neutral buoyancy. (2) The ceratites from the Germanic Basin had a nektonic mode of life. (3) The discoconic and platyconic forms were good at continuous swimming at low velocities and the oxyconic forms were faster swimmers, capable of rapid acceleration. (4) Germanic ceratites probably were good backward swimmers. (5) The orientation of the aperture of ceratites was between 608 and 908, depending on body chamber length and perhaps also on adult modifications. (6) Because of the more or less horizontal aperture, the steerage was probably good. (7) Ceratites with shells encrusted by bivalves syn vivo reacted (at least sometimes) with a change in coiling. (8) Sexual dimorphism among ceratites appears likely but needs further study. (9) Ceratites were rather stenohaline than euryhaline.

25

Acknowledgements Hans Hagdorn (Ingelfingen) put his collection of Muschelkalk ammonoids at our disposal and discussed several aspects of ceratite palaeobiology with us. Hans Hagdorn, Susan Turner (Kenmore, Queensland) and Nicolas Goudemand (Zu¨rich) thoughtfully proof-read the manuscript and offered substantial help in improving language as well as structure and content of the manuscript. Hans Rieber and Hugo Bucher (both Zu¨rich) contributed some important details and discussed some of our results with us. Ana Ma´rquez-Aliaga (Burjassot, Valencia) gave us some information on the Middle Triassic in Spain and Italy. Gu¨nter Schweigert (Stuttgart) kindly provided access to the substantial collections of the Staatliches Museum fu¨r Naturkunde Stuttgart. We appreciate the helpful work of the reviewers Franz T. Fu¨rsich (Wu¨rzburg), H. Keupp (Berlin) and Gerd E.G. Westermann (Hamilton, Ontario).

Appendix A. Data sets of all specimens used for the stratophenetic analysis. Data sets in bold type represent the largest specimen of each species in the material used

No. Tax. Strat. Inv. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

1 1 1 1 1 1 1 1 1 1 2 3 4 5 4 4 4 4 4 3 3 5

1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2

14745 F.L-6/94 25320/1 24507 24506 24503 24504 24528 24524 24514 14633 24662 v5/3831/1 25323/3 24570 24569 24575 24592 N178 24661 24666 24708

dm 50.6 53.6 71.7 65.4 59 60.7 67.9 66.9 50.7 56.2 69.7 85.3 60 65.6 83.1 66.3 69.7 56.3 86.2 84.5 58.4 66.8

ww wh

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa

nl nvl nl/nvl

17.6 15.1 22.3 29.3 19.1 18.8 19 21.1 17.6 17.6 24.7 36.1 18 21.6 27.1 24.4 23.1 17.8 29.1 30.1 28.5 19

7.1 11 12.8 10.6 13.2 11.5 11.3 11.3 9.5 9.9 11.2 16.6 10.7 15.7 14.7 14.3 13.9 11.4 20 19.2 12.2 15.8

2.42 1.86 1.94 2.26 2.21 2.18 2.18 2.27 2.09 2.04 2 1.9 2.33 2.12 2.24 2.08 2.14 2.13 2.02 1.93 1.99 2.28

11 13 11 11 9 10 13 13 12 13 13 13 10 9 10 10 10 10 12 13 8 9

26.1 24.9 23.9 33.8 27.5 30.1 34.4 32.8 25.9 29 34.2 43.2 29.1 29.5 42.8 31.4 36 26.6 25.6 41.8 28.2 29.7

0.35 0.28 0.31 0.45 0.32 0.31 0.28 0.32 0.35 0.31 0.35 0.42 0.33 0.33 0.33 0.37 0.33 0.32 0.34 0.36 0.49 0.28

0.52 0.46 0.33 0.52 0.47 0.50 0.51 0.49 0.51 0.52 0.49 0.51 0.49 0.45 0.52 0.47 0.52 0.47 0.30 0.49 0.48 0.44

0.14 0.21 0.18 0.16 0.22 0.19 0.17 0.17 0.19 0.18 0.16 0.19 0.18 0.24 0.18 0.22 0.20 0.20 0.23 0.23 0.21 0.24

0.31 0.43 0.15 0.35 0.30 0.35 0.36 0.31 0.40 0.40 0.40 0.43 0.29 0.31 0.36 0.35 0.39 0.33 0.35 0.43 0.68 0.24

1 1 1 1 1.5 1 1 1 1 1 1 1.5 1.5 2 1.5 1.5 1.5 1.5 2 2 2 2

1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1.5 1.5 1.5 2 2

18 12.7 18.6 14.3 12.9 14.7 15.3 14.6 12.2 12.7 18.6 22.8 15.4 15.6 20.9 15.4 17 12.2 23.4 22.8 18.1 15.6

19.2 14.5 19.9 15 16.4 16.5 16.6 16.2 14.1 14.5 21 24.5 17.3 18 21.7 17.3 18.1 14.4 26.1 24.5 21.5 18

1.07 1.14 1.07 1.05 1.27 1.12 1.08 1.11 1.16 1.14 1.13 1.07 1.12 1.15 1.04 1.12 1.06 1.18 1.12 1.07 1.19 1.15

23 19 21 19 19 18 22 20 22 19 18 19 20 22 17 20 20 20 21 19 18 22

0.48 0.68 0.52 0.58 0.47 0.56 0.59 0.65 0.55 0.68 0.72 0.68 0.50 0.41 0.59 0.50 0.50 0.50 0.57 0.68 0.44 0.41

(continued on next page)

26

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Appendix A (continued) No. Tax. Strat. Inv.

dm

ww wh

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa

nl nvl nl/nvl

23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76

70.6 72.6 84.4 87.3 81.9 59.6 76.1 71.3 73.7 65.7 70.5 64.2 63.1 78.9 65.1 69.3 65.2 59.9 79.7 79.3 100.2 87.6 76.8 74.7 65 102.7 74.1 65.7 76.6 110.7 85.8 76.8 98.3 108.7 87.5 69.4 103.1 87.4 82.8 88.7 78 100.2 110.6 70.1 96.3 74.6 79.3 80.2 68.2 77.7 72.8 79 98.8 79.6

20.5 20.3 25.4 24.9 22.3 17.6 26.6 22.6 20.5 18.8 19.6 24.3 16.8 17.7 23.4 24.6 22 16.3 17.8 26.4 34 26.7 20.3 29 19.1 31 25.3 19 19.6 31 26.3 27 26.6 30 26 21.6 29 31 24.1 30 20.5 28.7 33.7 22.5 27.3 21.8 24.7 23.4 16.9 24 21 22.1 28.8 23

16.1 17 19.9 26.1 16.4 12.6 15.6 14.6 19.4 15.2 20.9 16.9 16.5 21.6 15.3 10.7 10.1 14.6 19.5 22 30 25.5 23.4 21.5 18.4 27.5 18.8 18.5 19.5 39.1 28.3 22 37.3 42.3 23.8 19.5 24.9 22.9 21.5 31.4 22.4 30.8 27.7 29 36.6 25.2 20.6 23.4 20.8 23.8 17.3 25.5 31.5 26.8

2.24 2.08 1.78 1.88 1.78 1.99 1.99 2.03 1.78 1.99 1.81 2.3 1.19 1.89 2.07 2.53 2.05 1.96 2.04 2.09 1.95 1.79 1.75 1.72 1.79 1.76 1.72 1.91 1.87 1.87 1.74 1.77 1.84 1.73 1.99 2.1 2.2 1.82 2.03 2 1.88 1.77 1.84 1.97 1.75 1.83 1.96 1.89 1.91 1.88 1.94 1.98 1.84 1.97

10 10 9 10 12 14 10 12 10 11 9 9 10 14 9 9 7 10 10 9 8 5 7 11 9 7 10 9 10 10 9 10 9 10 11 12 12 11 10 11 15 12 8 12 10 10 9 11 12 10 11 11 9 10

5 8 6 6 2 1 2 2 8 8 8 8 9 9 8 2 2 6 9 10 10 7 7 7 12 7 10 11 11 13 13 11 15 15 16 16 16 24 16 16 16 17 20 16 17 18 19 17 16 19 19 16 19 19

2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6

24728 24702 24566 24562 24567 24565 24750 24749 33068 24747 24691 75207/2 25324/1 384 176 239 11588 24689 24685 24683 25326/3 25322/1 25728-1 Lu 65

75207/1 25281-1 25283-3 23075 10 3298 26944 24686 3831/5 25330/2 25332/4 25331/6

25297-1 25284-1

35.5 28.7 41 34.4 37.7 28.6 35.4 31.8 31.7 29.4 27.6 27.6 26.1 31.2 29.2 38 32.3 26.1 34.5 35.4 38.7 36.7 28.6 31.9 27.5 44.6 31.6 27.4 31.9 43.5 32.6 31.5 32.2 38.2 35.1 27.6 44.9 37.6 35.8 33.1 30.1 39 47.3 27.1 33.2 27.8 32.5 28.8 25.1 29.8 31.6 29 38.5 30.2

0.29 0.28 0.30 0.29 0.27 0.30 0.35 0.32 0.28 0.29 0.28 0.38 0.27 0.22 0.36 0.35 0.34 0.27 0.22 0.33 0.34 0.30 0.26 0.39 0.29 0.30 0.34 0.29 0.26 0.28 0.31 0.35 0.27 0.28 0.30 0.31 0.28 0.35 0.29 0.34 0.26 0.29 0.30 0.32 0.28 0.29 0.31 0.29 0.25 0.31 0.29 0.28 0.29 0.29

0.50 0.40 0.49 0.39 0.46 0.48 0.47 0.45 0.43 0.45 0.39 0.43 0.41 0.40 0.45 0.55 0.50 0.44 0.43 0.45 0.39 0.42 0.37 0.43 0.42 0.43 0.43 0.42 0.42 0.39 0.38 0.41 0.33 0.35 0.40 0.40 0.44 0.43 0.43 0.37 0.39 0.39 0.43 0.39 0.34 0.37 0.41 0.36 0.37 0.38 0.43 0.37 0.39 0.38

0.23 0.23 0.24 0.30 0.20 0.21 0.20 0.20 0.26 0.23 0.30 0.26 0.26 0.27 0.24 0.15 0.15 0.24 0.24 0.28 0.30 0.29 0.30 0.29 0.28 0.27 0.25 0.28 0.25 0.35 0.33 0.29 0.38 0.39 0.27 0.28 0.24 0.26 0.26 0.35 0.29 0.31 0.25 0.41 0.38 0.34 0.26 0.29 0.30 0.31 0.24 0.32 0.32 0.34

0.34 0.22 0.48 0.31 0.46 0.40 0.37 0.33 0.42 0.35 0.34 0.21 0.31 0.31 0.32 0.32 0.39 0.34 0.31 0.31 0.27 0.40 0.34 0.45 0.40 0.43 0.44 0.34 0.35 0.31 0.37 0.40 0.20 0.32 0.27 0.22 0.25 0.40 0.31 0.21 0.30 0.36 0.39 0.25 0.29 0.30 0.30 0.24 0.25 0.29 0.35 0.21 0.32 0.24

2 2 2 2 1 1 1 1 2 2 2 2 1.5 1.5 2 1 1 1.5 1.5 2 2 2 1 0.5 2 1 2 2 2 2 2 2 2.5 2.5 1.5 2 1.5 2 2 2 1.5 3.5 2 2 4 2.5 2 2.5 2 2 2.5 2.5 1.5 2

2 1 0 0.5 1 1 1.5 0.5 1 1.5 2 1.5 1 1.5 1 0.5 0 1.5 1 1 1.5 0 0.5 0.5 1 0 3 3 3 3 3 2.5 4 3.5 3 3 3 5 1.5 5 3.5 4 3 3 3 3 3 3 3 4 3 3 3 4

16.2 18.4 17.6 17.6 20.3 14.7 19 18.6 18.4 14.4 17.6 16 15.9 14.6 14.3 18.2 18 15.5 14.5 20 25.8 20.5 17.3 17.1 13.6 26.3 18 13.5 13.9 26 23 19 22.5 25.8 18 19 25.5 29 18.8 29 20.5 25.5 26 19 23.3 19.6 18.3 21 19 17.2 17.6 21 18.5 17

18.4 21.5 19.4 19.4 21 16.5 21 21 21.5 15.7 21.5 19.6 19 16.5 15.5 20 19.6 17.05 16.3 23 29.5 23.1 18.7 18.5 15.7 30.1 20.3 15.6 16.1 28.6 28.5 19.2 30.6 28.4 23 21.8 27.7 34 20.6 34 23.6 27.7 28.9 21.8 26.4 23.7 22.3 25 21.8 21 20.5 24 22.2 20

1.14 1.17 1.10 1.10 1.03 1.12 1.11 1.13 1.17 1.09 1.22 1.23 1.19 1.13 1.08 1.10 1.09 1.10 1.12 1.15 1.14 1.13 1.08 1.08 1.15 1.14 1.13 1.16 1.16 1.10 1.24 1.01 1.36 1.10 1.28 1.15 1.09 1.17 1.10 1.17 1.15 1.09 1.11 1.15 1.13 1.21 1.22 1.19 1.15 1.22 1.16 1.14 1.20 1.18

17 20 10 18 24 21 18 17 20 23 18 18 18 23 19 16 16 16 18 15 13 13 15 25 10 11 15 15 15 15 11 16 15 14 16 18 17 14 13 14 17 13 15 18 10 18 13 21 18 14 19 19 15 17

0.59 0.50 0.90 0.56 0.50 0.67 0.56 0.71 0.50 0.48 0.50 0.50 0.56 0.61 0.47 0.56 0.44 0.63 0.56 0.60 0.62 0.38 0.47 0.44 0.90 0.64 0.67 0.60 0.67 0.67 0.82 0.63 0.60 0.71 0.69 0.67 0.71 0.79 0.77 0.79 0.88 0.92 0.53 0.67 1.00 0.56 0.69 0.52 0.67 0.71 0.58 0.58 0.60 0.59

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

27

Appendix A (continued) No. Tax. Strat. Inv.

dm

ww wh

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa

nl nvl nl/nvl

77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129

89.7 95.9 91.7 117.5 80.7 97.7 81.5 86 83.6 89.8 120.7 96.8 87.3 88 87.5 77.8 86.3 103.1 101.6 78.1 62.9 70.8 67.2 64.6 86.3 131.3 102.1 119.3 80.1 88.8 100.5 149 148.5 112.4 116 128.1 77.9 90.7 96.7 95.2 127.3 107.2 90.4 96.1 90.7 121.8 78.3 83 82.5 119.7 121.4 119.7 116.2

24.6 33.5 26.9 35.2 24.6 26.6 25 30 29.9 27.9 37.8 29.1 24.6 21.4 24.3 24.9 26 36.1 41.7 28.5 20.3 24.7 25.5 18.6 22.9 52.3 41.3 46 29 31.9 41 53.4 57.7 46.3 48.7 52 28.5 32.5 28.3 28 43.1 37.4 30.1 28.1 31.8 42 28.6 30.5 30 35.2 44 41 43

26.4 33.7 30.5 40.9 24.3 33.4 24.9 24 23.7 33.7 40 32.2 21.8 29.1 28.9 30.5 31.6 31.8 35.6 20.8 15.7 20.3 20.2 17.9 22.4 35 25.1 31.4 19.5 20.3 29.3 44.9 43.9 31.3 39.2 34.4 16.8 23.7 29.4 27.1 33.2 28.7 25.7 25.5 24.5 43.9 22.7 22.7 25.8 39.2 24.6 27.2 28.5

1.89 1.89 1.86 2.17 1.89 1.84 1.83 2.12 2.11 1.83 1.89 1.79 1.96 1.69 1.87 1.77 1.61 1.69 1.75 2.08 1.83 1.47 2.35 1.94 1.56 2.1 2 1.99 2.04 2 1.66 1.99 2.34 2.1 2.39 1.9 2.17 2.12 1.8 2.34 2.09 2.11 2.05 1.47 1.92 1.86 2.03 2.08 1.91 2.01 1.99 1.99 2.2

11 12 13 11 12 12 11 10 12 13 15 10 9 11 11 12 12 10 11 10 11 10 11 10 10 11 10 9 12 10 10 11 10 11 10 10 10 10 10 11 10 10 10 11 10 11 12 13 10 12 10 11 11

16 21 21 21 23 22 25 24 24 22 23 30 23 22 23 23 21 23 25 27 28 28 28 29 30 31 31 31 31 31 31 35 35 33 33 31 31 31 31 31 32 31 32 31 31 32 32 32 32 32 33 31 33

6 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 8 8 9 9 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8

25285-1

75284 17993/30

166 75210/3 25333/3 25333/5 25334/1 59

19/58 9771

720

21082

36.2 36.8 23.1 40.2 31.4 35.1 32 33 35.3 14.1 46.9 35.4 36.3 32.6 32.5 26.8 31.3 40 24.8 31.4 28.7 29.3 31.2 26.3 37.2 48.5 44.3 46.7 35.8 39.7 38.5 57.6 63.5 51.6 54.6 52.5 34.9 37.2 38.2 40.3 58.6 44.3 37.6 40.1 37.6 44.7 30.8 35.9 31.8 45.2 50 32 49.1

0.27 0.35 0.29 0.30 0.30 0.27 0.31 0.35 0.36 0.31 0.31 0.30 0.28 0.24 0.28 0.32 0.30 0.35 0.41 0.36 0.32 0.35 0.38 0.29 0.27 0.40 0.40 0.39 0.36 0.36 0.41 0.36 0.39 0.41 0.42 0.41 0.37 0.36 0.29 0.29 0.34 0.35 0.33 0.29 0.35 0.34 0.37 0.37 0.36 0.29 0.36 0.34 0.37

0.40 0.38 0.25 0.34 0.39 0.36 0.39 0.38 0.42 0.16 0.39 0.37 0.42 0.37 0.37 0.34 0.36 0.39 0.24 0.40 0.46 0.41 0.46 0.41 0.43 0.37 0.43 0.39 0.45 0.45 0.38 0.39 0.43 0.46 0.47 0.41 0.45 0.41 0.40 0.42 0.46 0.41 0.42 0.42 0.41 0.37 0.39 0.43 0.39 0.38 0.41 0.27 0.42

0.29 0.35 0.33 0.35 0.30 0.34 0.31 0.28 0.28 0.38 0.33 0.33 0.25 0.33 0.33 0.39 0.37 0.31 0.35 0.27 0.25 0.29 0.30 0.28 0.26 0.27 0.25 0.26 0.24 0.23 0.29 0.30 0.30 0.28 0.34 0.27 0.22 0.26 0.30 0.28 0.26 0.27 0.28 0.27 0.27 0.36 0.29 0.27 0.31 0.33 0.20 0.23 0.25

0.33 0.29 0.25 0.06 0.30 0.27 0.34 0.18 0.26 0.31 0.30 0.31 0.31 0.37 0.28 0.28 0.42 0.40 0.24 0.43 0.57 0.25 0.31 0.54 0.16 0.33 0.26 0.33 0.35 0.41 0.25 0.19 0.33 0.25 0.33 0.28 0.24 0.36 0.18 0.33 0.25 0.27 0.58 0.33 0.27 0.24 0.29 0.28 0.22 0.29 0.25 0.23

2 4 4 4 2 4 2 2 2 4 2 2 2 2.5 4 2 4 4 2 2 2 2 2 2 2 5 4.5 2 2 2 2 5 5 5 5 5 2 2 5 5 2 5 2 2 2 3.4 2 2 2 2 5 5 2.5

3.5 4 4 4 4 4 4 5 5 4 4 5 2.5 4 4 3.5 4 5 5 2.5 2 2 2 2.5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 4.5 5 5 5 5 5 5 5 5 5 5 5 5

18 33.1 24.7 29.6 18.7 21.7 18.7 29 28.7 28.2 32.6 35 21.7 19 23 18.8 24 35 25 21.3 16.7 23 26 17 23 44 38 47 19 24 38 50 45 54 48 33 24.4 22.7 22.8 22.6 32.4 31.6 22.6 22.7 22 33 19.7 23.3 23.1 32.3 34 32.4 53

24 37.9 29.3 35.1 22.4 24.9 22.4 34 33.6 32.2 35.2 47 23 22 26.7 22.5 29 47 33 26.6 20.6 27 30.8 19.5 26.7 55 49 54 21 30 48 65 60 61 54 44 30.1 27.5 28 27 42.2 40.5 27.5 27 26.9 43 23.4 29.8 29.5 42.8 40 42.6 60

1.33 1.15 1.19 1.19 1.20 1.15 1.20 1.17 1.17 1.14 1.08 1.34 1.06 1.16 1.16 1.20 1.21 1.34 1.32 1.25 1.23 1.17 1.18 1.15 1.16 1.25 1.29 1.15 1.11 1.25 1.26 1.30 1.33 1.13 1.13 1.33 1.23 1.21 1.23 1.19 1.30 1.28 1.22 1.19 1.22 1.30 1.19 1.28 1.28 1.33 1.18 1.31 1.13

16 12 19 11 16 15 12 13 15 14 18 12 17 16 11 16 17 12 13 15 20 16 17 18 12 11 10 10 18 13 13 11 10 12 10 11 12 14 12 13 12 12 14 14 13 11 15 17 12 13 11 13 12

0.69 1.00 0.68 1.00 0.75 0.80 0.92 0.77 0.80 0.93 0.83 0.83 0.53 0.69 1.00 0.75 0.71 0.83 0.85 0.67 0.55 0.63 0.65 0.56 0.83 1.00 1.00 0.90 0.67 0.77 0.77 1.00 1.00 0.92 1.00 0.91 0.83 0.71 0.83 0.85 0.83 0.83 0.71 0.79 0.77 1.00 0.80 0.76 0.83 0.92 0.91 0.85 0.92

(continued on next page)

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C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Appendix A (continued) No. Tax. Strat. Inv.

dm

ww wh

130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183

117.3 100.5 90.4 96 108.1 81.9 90.3 85.7 135 106.6 100.9 98.5 110.2 103.2 77.3 100.6 92.4 134.5 87.5 116.4 115 81.5 107.6 101.6 128.3 141 72.7 75 94.3 164.4 120.6 132.3 199.4 178 155.9 159.8 129.6 123.7 195 129.6 102.3 160 169 116 110.1 115 157 173 234 141.1 114.5 88.3 116.3 116.4

44 50.5 31.7 2.08 37 42 23.6 2.16 34 35.9 23.6 2.29 35 37 35 1.64 40.5 47 28.4 1.88 25.4 34.7 23.8 1.97 28.8 35.5 24.8 1.99 28.5 35.6 24.8 2.11 44.5 56.8 38.6 2.1 35.8 44.7 28.7 2.04 34.4 43.3 26.9 2 36.8 44.1 28.8 2.3 44.2 46.2 30.8 1.88 37.4 44.1 26.6 2.06 28.2 33.3 18.9 2.02 36.7 47.1 21.1 2.11 29.6 40.8 23.5 2.05 51.7 57 38.3 2.03 29 33.8 27.1 1.78 42.6 46.9 33.3 2 42.3 48.5 34.3 2.08 30.3 35.8 20.7 2.1 44.3 46 29.1 2.26 35 39.4 31.1 2.08 54.5 57.5 36 2.14 50.2 60 38.5 2.01 26.4 33.2 16.1 2.17 27.9 33.6 17.4 2.19 35.9 41.8 23.4 2.22 55 64.8 58.8 2.03 38 55.9 28.1 1.53 49 58.1 39.6 2.21 71.2 85.1 46.6 2.14 69 68.5 59.4 1.75 57.7 72.2 39.9 2.3 59 70.3 38.3 2.17 48.5 59.4 30.6 2.14 48.6 54.6 31.9 2.09 70 77.6 63.3 1.98 48.3 53.9 39.6 2.09 34.4 47.2 22.1 2.06 58.7 71.6 38.9 2.04 65 70 50.1 1.95 41 52.3 25.6 1.98 39 46 29.6 1.9 40.4 47.2 26.1 0.02 58 61.2 49.9 2 67.7 82.3 41.6 2.11 81 113.6 66.6 2.34 46.5 54.6 46.5 2.04 26.9 41.9 32.5 2.01 27.1 36.4 23.6 1.87 28 48.5 28.2 1.94 32 51.2 27.2 2.01

33 33 33 34 31 26 31 33 31 31 31 31 31 31 31 31 31 33 26 31 31 31 31 33 33 31 30 31 31 35 36 35 35 35 35 35 35 34 35 34 36 35 35 36 34 36 35 36 35 34 38 37 37 37

8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 10 10 10 10

52 14637 16505 26940-15 41 58 54 39 52 43 37 50 25255/8 25255/9 25255/7 25255/24 42 25255/28 25255/29 25255/12 26940/3 26940/14 55 21085

24500

46 24442 24439 21849 14436 25338/1 393 397

75283 9735 75200

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa 0.38 0.37 0.38 0.36 0.37 0.31 0.32 0.33 0.33 0.34 0.34 0.37 0.40 0.36 0.36 0.36 0.32 0.38 0.33 0.37 0.37 0.37 0.41 0.34 0.42 0.36 0.36 0.37 0.38 0.33 0.32 0.37 0.36 0.39 0.37 0.37 0.37 0.39 0.36 0.37 0.34 0.37 0.38 0.35 0.35 0.35 0.37 0.39 0.35 0.33 0.23 0.31 0.24 0.27

0.43 0.42 0.40 0.39 0.43 0.42 0.39 0.42 0.42 0.42 0.43 0.45 0.42 0.43 0.43 0.47 0.44 0.42 0.39 0.40 0.42 0.44 0.43 0.39 0.45 0.43 0.46 0.45 0.44 0.39 0.46 0.44 0.43 0.38 0.46 0.44 0.46 0.44 0.40 0.42 0.46 0.45 0.41 0.45 0.42 0.41 0.39 0.48 0.49 0.39 0.37 0.41 0.42 0.44

0.27 0.23 0.26 0.36 0.26 0.29 0.27 0.29 0.29 0.27 0.27 0.29 0.28 0.26 0.24 0.21 0.25 0.28 0.31 0.29 0.30 0.25 0.27 0.31 0.28 0.27 0.22 0.23 0.25 0.36 0.23 0.30 0.23 0.33 0.26 0.24 0.24 0.26 0.32 0.31 0.22 0.24 0.30 0.22 0.27 2.27 0.32 0.24 0.28 0.33 0.28 0.27 0.24 0.23

0.29 0.24 0.14 0.43 0.38 0.32 0.26 0.25 0.26 0.28 0.32 0.24 0.35 0.29 0.31 0.33 0.32 0.30 0.35 0.28 0.27 0.30 0.22 0.21 0.30 0.31 0.30 0.28 0.26 0.24 0.58 0.25 0.26 0.37 0.27 0.27 0.31 0.30 0.27 0.26 0.34 0.33 0.31 0.36 0.34 2.39 0.25 0.34 0.29 0.23 0.20 0.35 0.32 0.33

5 5 2 2 5 2 2 2 2 2 2 2 2.5 2 1.5 2 2 5 2 4 5 2 2.5 4 2.5 5 2 2 2 2 5 5 5 5 5 5 2 5 5 2 2 2 5 1.5 2.5 2 5 5 5 5 2 0 0 0

5 5 5 5 5 2 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 3 5 5 5 5 5 5 5 5 5 5 5 0 0 0 0

33 27 28.5 27.3 31.6 19.8 24.4 22.8 40.5 31.6 25.5 27.3 40 23.8 19.7 27 22.7 45 22.8 32.3 36 23.3 40 33.5 35 38 19.7 20 27.3 46.7 32 49 71.2 46.7 46 47 49 44 72 44 31 46.9 46.7 32 30.1 31 46 51 58 38 22 27.1 28 27.3

42 36 38 40.6 40.5 21.6 30.1 29 51.5 40.5 35.4 40.6 49.9 30.7 23.4 36 27.5 57 29 42.8 43.1 29.8 49.9 42.9 41 45 23.4 24 40.6 55.2 45 55 82 55.2 53 53.7 55 52 82 53 65 54 55.2 40 41 44 53 57 64 45 27 27.1 28 27.3

1.27 1.33 1.33 1.49 1.28 1.09 1.23 1.27 1.27 1.28 1.39 1.49 1.25 1.29 1.19 1.33 1.21 1.27 1.27 1.33 1.20 1.28 1.25 1.28 1.17 1.18 1.19 1.20 1.49 1.18 1.41 1.12 1.15 1.18 1.15 1.14 1.12 1.18 1.14 1.20 2.10 1.15 1.18 1.25 1.36 1.42 1.15 1.12 1.10 1.18 1.23 1.00 1.00 1.00

nl nvl nl/nvl 14 12 11 8 10 12 10 9 10 10 10 8 12 11 12 12 10 11 9 11 11 9 12 11 10 10 12 10 11 12 10 13 14 14 12 12 13 13 13 14 9 16 10 11 13 12 13 13 15 13 9 0 0 0

15 14 13 9 12 18 12 14 13 12 11 9 14 13 15 14 14 13 14 13 11 12 14 13 11 10 15 13 12 14 12 14 16 14 13 12 18 14 13 16 11 17 11 11 18 14 13 15 15 13 11 14 0 0

0.93 0.86 0.85 0.89 0.83 0.67 0.83 0.64 0.77 0.83 0.91 0.89 0.86 0.85 0.80 0.86 0.71 0.85 0.64 0.85 1.00 0.75 0.86 0.85 0.91 1.00 0.80 0.77 0.92 0.86 0.83 0.93 0.88 1.00 0.92 1.00 0.72 0.93 1.00 0.88 0.82 0.94 0.91 1.00 0.72 0.86 1.00 0.87 1.00 1.00 0.82 0.00 0.50 0.50

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

29

Appendix A (continued) No. Tax. Strat. Inv. 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236

38 38 39 39 40 40 40 39 40 39 40 39 40 42 42 41 46 43 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 44 47 44 44 47 44 46 45 45 48 48 48 48 48 48 48

10 10 11 11 11 11 11 11 11 11 11 11 11 12 12 12 12 12 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 14 14 14 14 14 14 14

203 21075

dm

126.1 128.1 119.7 105.2 96.4 18717 91.6 97.7 255 103.1 238 63.4 416 71.9 105.4 115.2 125.2 26946 99.6 113.3 86.6 25418-1 104.8 25416 144 20324 196 24950 175 22972 163 183 24953 192 24953/952 189 24937 181 210 6824/1 202 16714 243 233 24941 185 232 24931 186 24934 165 24938 223 20 226 24936 153 172 196 963.965 188 25426 171 25426 155 25425-1 190 747 188 118 148 378 177 25424 189 13581 149 224 25408/2 146 25305 161 25468-4 182 14659 290

ww wh

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa

nl nvl nl/nvl

32 35 31 33 29 30.6 31 35.3 20.1 22.8 27.2 30 34 42.7 53.2 30.3 39 57 78.8 63.4 71 66.8 76.1 76.9 68.8 79 74 92.8 90 73 90.8 72 71.7 79 80.5 61.7 75 77 69.9 63 61.5 70 85.7 42.5 56 67 58 46.3 75.4 45 50.8 57 98

44.4 37.3 29.7 32.1 19.8 27.1 22.2 26 12.4 16.6 30.3 29.3 39.4 22.8 22.8 21.4 22.6 44.8 54.8 52.4 37.5 50.6 51.4 51.6 47.1 61.4 53.3 54.8 55.1 43.1 52.2 52.3 73.2 51.2 52.9 34.5 48.6 57.1 47.6 43.2 35.9 44.8 49.6 27.1 33 46 44.1 22.4 54.4 30.8 28.8 42.6 58.6

1.92 1.81 1.96 1.61 1.9 1.81 1.78 2 1.94 2.05 1.82 2.2 1.92 2 1.89 1.95 2.38 1.94 1.93 1.97 2.2 2.18 1.91 1.82 2.02 1.99 2.24 2.43 2.32 2.26 2.05 2.21 1.85 2.21 2.07 2.33 2.05 2.02 1.91 2.06 2.26 1.92 2.23 2.24 1.86 1.77 1.84 2.26 1.84 2.29 2.09 1.96 1.94

10 8 9 10 11 11 13 12 5 8 11 12 12 11 13 11 14 11 12 12 15 12 11 12 12 12 10 10 11 11 11 11 10 12 11 13 11 11 12 15 11 12 14 13 17 12 10 11 11 9 11 13 11

46.1 49.1 50.2 42 40.3 35.2 40 44.3 29 31.1 41.8 49.8 49.5 46.7 51.6 37.5 50.7 61 87.8 68.7 72.4 85.3 86.9 92.9 83.2 87.9 85.7 127.3 111.1 86.7 106.5 72.1 74.4 116.1 107.4 67.2 64.3 82.6 90 72.4 69.7 81 81.4 51.8 72.2 82 54.8 75 103 68 76.4 75 139.4

0.25 0.27 0.26 0.31 0.30 0.33 0.32 0.34 0.32 0.32 0.26 0.26 0.27 0.43 0.47 0.35 0.37 0.40 0.40 0.36 0.44 0.37 0.40 0.41 0.38 0.38 0.37 0.38 0.39 0.39 0.39 0.39 0.43 0.35 0.36 0.40 0.44 0.39 0.37 0.37 0.40 0.37 0.46 0.36 0.38 0.38 0.31 0.31 0.34 0.31 0.32 0.31 0.34

0.37 0.38 0.42 0.40 0.42 0.38 0.41 0.43 0.46 0.43 0.40 0.43 0.40 0.47 0.46 0.43 0.48 0.42 0.45 0.39 0.44 0.47 0.45 0.49 0.46 0.42 0.42 0.52 0.48 0.47 0.46 0.39 0.45 0.52 0.48 0.44 0.37 0.42 0.48 0.42 0.45 0.43 0.43 0.44 0.49 0.46 0.29 0.50 0.46 0.47 0.47 0.41 0.48

0.35 0.29 0.25 0.31 0.21 0.30 0.23 0.25 0.20 0.23 0.29 0.25 0.31 0.23 0.20 0.25 0.22 0.31 0.28 0.30 0.23 0.28 0.27 0.27 0.26 0.29 0.26 0.23 0.24 0.23 0.23 0.28 0.44 0.23 0.23 0.23 0.28 0.29 0.25 0.25 0.23 0.24 0.26 0.23 0.22 0.26 0.23 0.15 0.24 0.21 0.18 0.23 0.20

0.24 0.33 0.32 0.47 0.34 0.33 0.39 0.32 0.38 0.30 0.35 0.24 0.30 0.37 0.40 0.34 0.27 0.34 0.37 0.27 0.27 0.31 0.39 0.47 0.35 0.31 0.22 0.32 0.28 0.28 0.34 0.15 0.41 0.37 0.36 0.21 0.19 0.30 0.42 0.28 0.25 0.35 0.24 0.25 0.26 0.25 0.32 0.34 0.43 0.27 0.35 0.31 0.41

0.5 2 2 2.5 1 1 1 2 1 2 2 2 2.5 1.5 1 1.5 2 4 3 2 2 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 3 2 2 1 4 2 2.5 5 1 0.5 1 0.5 1 1.5 1.5

0 0 3.5 3 3 3.5 3 4 3 3.5 4 4 3.5 5 5 5 3 5 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 3 4.5 4 3 4 3 5 5 3.5 4 4 4 4 4 4

27.9 35 32.4 26 27 26.8 27.5 26.7 20.6 21.8 27 32.4 30.7 31.7 41.4 47 31.3 37 71.4 57 55 64 63 63 65 64 67 94 85 66 93 65 55 70 71 62 62 67 57 57 62 59 77 32 37.3 63 44 37 65 36 39.2 43 83

34.4 35 35.6 29 28 27.9 29 28.3 23 25.2 30 35.6 35.4 35.8 55.5 65 33 42 88.8 68 69 82 77 76 83 78 80 105 97 78 101 82 69 80 81 71 75 79 65 68 71 66 93 35 41.6 75 50 40 72 39.5 40.5 48 91

1.23 1.00 1.10 1.12 1.04 1.04 1.05 1.06 1.12 1.16 1.11 1.10 1.15 1.13 1.34 1.38 1.05 1.14 1.24 1.19 1.25 1.28 1.22 1.21 1.28 1.22 1.19 1.12 1.14 1.18 1.09 1.26 1.25 1.14 1.14 1.15 1.21 1.18 1.14 1.19 1.15 1.12 1.21 1.09 1.12 1.19 1.14 1.08 1.11 1.10 1.03 1.12 1.10

12 11 12 14 14 15 15 16 21 16 15 18 16 17 18 17 17 12 12 13 19 13 11 12 12 13 11 10 11 12 11 11 10 12 12 15 12 11 13 16 17 12 15 16 20 12 19 20 19 18 22 20 15

0.83 0.73 0.75 0.71 0.79 0.73 0.87 0.75 0.24 0.50 0.73 0.67 0.75 0.65 0.72 0.65 0.82 0.92 1.00 0.92 0.79 0.92 1.00 1.00 1.00 0.92 0.91 1.00 1.00 0.92 1.00 1.00 1.00 1.00 0.92 0.87 0.92 1.00 0.92 0.94 0.65 1.00 0.93 0.81 0.85 1.00 0.53 0.55 0.58 0.50 0.50 0.65 0.73

(continued on next page)

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C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Appendix A (continued) No. Tax. Strat. Inv.

dm

ww wh

uw

WER ww/dm wh/dm uw/dm IZR sph sbc samin samax sa

nl nvl nl/nvl

237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275

178 249 244 246 272 165 173 150 201 172 212 198 130.2 250 164 152 160 252 255 179 198 248 161 257 174 233 222 250 203 190 267 335 285 151 169 378 355 295 304

59.8 69 68 62 66.8 39 41 39.1 47.9 43 51 69.6 37.2 78 61 59 40 74 80 63 70 54 37 60 49 49 47 49 52 50.5 61 80 65 39 40 91 82 67 71

25.9 54.1 56.5 51 62.4 26.4 28.3 21.4 38 27.1 42.4 43.4 15.5 44.1 30.8 33.7 28.9 47.6 50.8 28.8 39.8 42 22.4 65.1 36.3 26.3 25.8 28.6 24.5 40.3 40.9 80 68.5 22.7 37.8 84.4 80.4 57 67.4

2.19 2.1 2.01 2 2.23 2.25 1.95 2.24 2.39 2.11 2.23 2.09 2.27 2.24 2.35 2.2 2 2 2.1 2.12 2.06 2.1 2.17 2.11 2.21 2.02 2.7 2.3 2.29 2.04 2.2 3.6 1.9 2.58 2.14 1.88 1.85 2.09 1.75

14 14 0 0 11 11 0 10 13 0 12 14 0 14 14 13 12 13 13 11 12 0 11 14 13 0 0 0 0 15 0 11 12 13 13 15 13 0 12

49 50 51 51 50 51 51 51 50 51 50 49 51 49 49 49 51 51 51 50 50 51 51 52 52 53 53 53 53 52 53 52 52 53 52 52 52 53 52

15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 16 16 16 16 16 16 16 16 16 16 16 16 16 16 16 16

18995-2 11615 16004

25400/4 25303 24498 24496 25422-1 25411-9 25411-2 25411-3 25469-4 25409-2 25409-9 25421-4 25422-6 25409-10 18995-3 25380 25377 21080/1 21080/2 25391 8950/2 27600 36 45 25401-1 25397-4 25394-4 25394-1 25397-3

83.7 115 106.3 104.6 123.5 89 89.1 76.1 98.6 85.4 104.5 86.8 68.8 117.5 78 68 73.4 119 121.4 94.3 85.2 122 82.3 107.4 84.1 121.7 119.3 123.9 106 85.5 132 140 127.8 82.7 77.6 162 146 138 136.1

0.34 0.28 0.28 0.25 0.25 0.24 0.24 0.26 0.24 0.25 0.24 0.35 0.29 0.31 0.37 0.39 0.25 0.29 0.31 0.35 0.35 0.22 0.23 0.23 0.28 0.21 0.21 0.19 0.26 0.27 0.23 0.24 0.23 0.26 0.24 0.24 0.23 0.23 0.23

0.47 0.46 0.44 0.43 0.45 0.54 0.52 0.51 0.49 0.50 0.49 0.44 0.53 0.47 0.48 0.45 0.46 0.47 0.48 0.53 0.43 0.49 0.51 0.42 0.48 0.52 0.54 0.50 0.52 0.45 0.49 0.42 0.45 0.55 0.46 0.43 0.41 0.47 0.45

0.15 0.22 0.23 0.21 0.23 0.16 0.16 0.14 0.19 0.16 0.20 0.22 0.12 0.18 0.19 0.22 0.18 0.19 0.20 0.16 0.20 0.17 0.14 0.25 0.21 0.11 0.12 0.11 0.12 0.21 0.15 0.24 0.24 0.15 0.22 0.22 0.23 0.19 0.22

0.31 0.33 0.32 0.31 0.27 0.38 0.45 0.35 0.28 0.37 0.33 0.30 0.36 0.29 0.27 0.27 0.36 0.38 0.35 0.41 0.30 0.37 0.36 0.26 0.32 0.43 0.27 0.31 0.35 0.33 0.34 0.31 0.39 0.31 0.31 0.37 0.36 0.34 0.46

2 3 0 0 1 1 0 1 1 1 0 2 0 2 2 2 1 1.5 3 1.5 1 0 1 1 1 0 0 0 0 1 0 1 1 1 1 1 1 0 2

4 3.5 0 0 3.5 0 0 1 3 0 0 4 0 4 4 4 0 4 3.5 3.5 3.5 0 0 1 1 0 0 0 0 1 0 1 1 0 1 3 3 0 0.5

42 59 68 51 57 32 33 29 45.4 35 45 57 37.2 61 42 40 35 48.6 55 32.7 57 45 33 51 43 42 40 41 44 49.5 54 68 58 28.4 34 79 73 60 62

48 65 68 51 65 32 33 30 51 35 49 64 37.2 69 48 45.3 37 51 62 35 64 45 33 55 47 42 40 41 44 52 54 75 63 29 37 87 80 65 67

1.14 1.10 1.00 1.00 1.14 1.00 1.00 1.03 1.12 1.00 1.09 1.12 1.00 1.13 1.14 1.13 1.06 1.05 1.13 1.07 1.12 1.00 1.00 1.08 1.09 1.00 1.00 1.00 1.00 1.05 1.00 1.10 1.09 1.02 1.09 1.10 1.10 1.08 1.08

17 18 0 0 18 22 0 20 21 0 17 18 0 18 17 17 18 20 22 12 17 0 19 30 24 0 0 0 0 27 0 12 18 36 25 20 18 0 17

0.82 0.78 0.50 0.50 0.61 0.50 0.50 0.50 0.62 0.50 0.71 0.78 0.50 0.78 0.82 0.76 0.67 0.65 0.59 0.92 0.71 0.50 0.58 0.47 0.54 0.50 0.50 0.50 0.50 0.56 0.50 0.92 0.67 0.36 0.52 0.75 0.72 0.50 0.71

Tax: taxa code (see Appendix B); strat: stratigraphy code (see text); inv: inventory number of the Staatliches Museum fu¨r Naturkunde Stuttgart (SMNS).

C. Klug et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 221 (2005) 7–34

Appendix B. Taxa code table (dtaxT in Appendix A) Abbreviations: P., Paraceratites; C., Ceratites; D., Discoceratites.

Species

Tax

P. atavus atavus P. atavus discus P. flexuosus bussei P. flexuosus flexuosus C. primitivus P. philippii neolaevis P. philippii philippii C. pulcher pulcher C. pulcher angustus C. robustus robustus C. robustus transgressor C. robustus stolleyi C. raricostatus C. transgressor C. rarinodosus C. compressus compressus C. compressus similis C. compressus crassior C. compressus apertus C. distractus C. evolutus evolutus C. evolutus tenuis C. evolutus subspinosus C. evolutus praecursor C. evolutus bispinatus C. armatus riedeli C. armatus nobilis C. armatus muensteri C. armatus posseckeri C. spinosus praespinosus C. spinosus spinosus C. spinosus capricornu C. spinosus obesus C. spinosus multicostatus C. spinosus penndorfi C. spinosus postspinosus C. enodis C. laevigatus C. sublaevigatus C. hercynus C. praenodosus C. macrocephalus C. bivolutus C. nodosus nodosus C. optimus D. parvus D. laevis D. weyeri

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48

31

Appendix B (continued) Species

Tax

D. D. D. D. D.

49 50 51 52 53

levalloisii alticella diversus dorsoplanus meissnerianus semipartitus

Appendix C. Structure matrix, eigenvalues of the canonical discriminant function analysis using the complete data set (missing values interpolated)

Structure matrix Function 1 sbc sph sa ww/dm wer wh/dm uw/dm izr nl/nvl

2 0.964T 0.342T 0.269T 0.476 0.017 0.187 0.055 0.078 0.521

3 0.192 0.021 0.158 0.751T 0.222T 0.573 0.121 0.057 0.185

4 0.137 0.131 0.023 0.343 0.144 0.788T 0.136T 0.089T .042

0.125 0.269 0.033 0.304 0.007 0.121 0.033 0.028 0.832T

Pooled within-groups correlations between discriminating variables and standardised canonical discriminant functions. Variables ordered by absolute size of correlation within function. T Largest absolute correlation between each variable and any discriminant function.

Eigenvalues Function

Eigenvalue

% of variance

Cumulative %

Canonical correlation

1 2 3 4

3.825a 1.012a 0.441a 0.292a

68.7 18.2 7.9 5.2

68.7 86.8 94.8 100.0

0.890 0.709 0.553 0.475

a

First 4 canonical discriminant functions were used in the analysis.

References Aigner, T., 1985. Storm depositional systems. Lecture Notes in Earth Sciences 3, 1 – 174. Aigner, T., Bachmann, G.H., 1992. Sequence-stratigraphic framework of the German Triassic. Sedimentary Geology 80, 115 – 135.

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