Dienerian (Early Triassic) ammonoids from the Northern Indian Margin

by David Ware, Hugo Bucher, Thomas Brühwiler,
Elke Schneebeli-Hermann, Peter A. Hochuli†, Leopold Krystyn,
Ghazala Roohi, Khalil Ur-Rehman and Amir Yaseen

Acknowledgement
Financial support for the publication of this issue of
Fossils and Strata was provided by the Lethaia Foundation

Foreword Dienerian (Early Triassic) ammonoids and the Early Triassic biotic recovery: a review
1. Present work
2. Definitions of stages and substages
3. Biochronology: the Unitary Association Method and terminology

Griesbachian and Dienerian (Early Triassic) ammonoids from the Salt Range, Pakistan

Griesbachian and Dienerian (Early Triassic) ammonoids from the Salt Range, Pakistan
1. Introduction
2. Geological framework
3. Biostratigraphy
4. Conclusion
5. Systematic palaeontology
  By David Ware and Hugo Bucher
6. References

Dienerian (Early Triassic) ammonoids from Spiti, Himachal Pradesh, India

Dienerian (Early Triassic) ammonoids from Spiti, Himachal Pradesh, India
1. Introduction
2. Geological framework
3. Biostratigraphy
4. Conclusion
5. Systematic palaeontology
6. References

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List of Illustrations

Foreword

Fig. 1. Total generic richness (black bold line, all ammonoids; grey lines, major ammonoid groups) and mean Chao2 estimate of the overall generic richness with its 95% confidence interval (large circles with vertical bars). The Early Triassic is highlighted in dark grey. PTB, Permian–Triassic boundary; 1, Kasimovian; 2, Gzhelian; 3, Asselian; 4, Sakmarian; 5, Artinskian; 6, Kungurian; 7, Roadian; 8, Wordian; 9, Capitanian; 10, Wuchiapingian; unlabelled successive intervals, Changhsingian, Griesbachian, Dienerian, Smithian; 15, Spathian; 16, Early Anisian; 17, Middle Anisian; 18, Late Anisian; 19, Ladinian; 20, Early Carnian; 21, Late Carnian; 22, Early Norian; 23, Middle Norian; 24, Late Norian; 25, Rhaetian. Modified after Brayard et al. (2009).
Fig. 2. Lower Triassic stage and substage subdivision (Ogg 2012) calibrated with recently published radiometric ages from South China.
Fig. 3. Simplified palaeogeographical map of the Early Triassic with the palaeopositions of the studied localities (white stars) and of other localities mentioned in the text (black stars). Modified after Brayard et al. (2006).

Chapter 1

Fig. 1. A. Palaeogeographical map of the Early Triassic with the palaeoposition of the Salt Range (modified from Brayard et al. 2006). B. Map of Pakistan with position of the studied area (black star). C. Location map of sampled sections in the Salt Range (modified after Brühwiler et al. 2012).
Fig. 2. Polished section of a bed belonging to the Ambites lilangensis Regional Zone (Nam100 and equivalents) from Nammal Nala, showing the typical facies of limestone beds at the base of the Ceratite Marls, with an accumulation of imbricated shells at its base with a nautiloid (N), several imbricated complete and broken ammonoids (A), and bioturbation increasing towards the top of the bed. Natural size, arrow indicates top of bed. Specimen PIMUZ30235.
Fig. 3. Section near the entrance of Nammal Nala (N32°39ʹ27.6″, E71°47ʹ29.2″). KM.d: Kathwai Member, dolomitic unit; KM.l: Kathwai Member, limestone unit; LCL: Lower Ceratite Limestone; CM: Ceratite Marls. Note the presence of small normal faults (black lines).
Fig. 4. Composite section of Nammal Nala: lithostratigraphy, ammonoid occurrences and biostratigraphy. Faunal content of NAM543, NAM718 and NAM540: see Brühwiler et al. (2012).
Fig. 5. Section above the village of Chiddru (N32°32ʹ59.7″, E71°47ʹ55.9″) on the West side of the gorge, previously described by Kummel & Teichert (1966, 1970) and Kummel (1966). Hammer highlighted for scale. Chiddru Fm., Chiddru formation; KM, Kathwai Member; LCL, Lower Ceratite Limestone; CM, Ceratite Marls.
Fig. 6. Composite section of Chiddru: lithostratigraphy, ammonoid occurrences and biostratigraphy. Faunal content of CHI63, CHI52, CHI104 and CHI53: see Brühwiler et al. (2012).
Fig. 7. Section about 1 km south-east of the village of Amb (N32°29ʹ48.1″, E71°56ʹ20.6″). Scale indicated by a geologist (circled) in the middle of the view. KM.d: Kathwai Member, dolomitic unit; KM.l: Kathwai Member, limestone unit; LCL: Lower Ceratite Limestone; CM: Ceratite Marls.
Fig. 8. Composite section of Amb: lithostratigraphy, ammonoid occurrences and biostratigraphy. AMB53 and AMB11 were collected on a dipslope formed by the two uppermost beds of the Lower Ceratite Limestone, without any further distinction.
Fig. 9. Section of Munta Nala, about 2.5 km West of the village of Wargal (N32°27ʹ07″, E72°01ʹ56.7″; Kummel 1966; Kummel & Teichert 1966, 1970,). Thickness of Kathwai Member and Lower Ceratite Limestone amounts to ca. 6 m. KM.d: Kathwai Member, dolomitic unit; KM.l: Kathwai Member, limestone unit; LCL: Lower Ceratite Limestone; CM: Ceratite Marls.
Fig. 10. Composite section of Wargal: lithostratigraphy, ammonoid occurrences and biostratigraphy. WAR11 and WAR100 were collected on a dipslope formed by several beds in the middle part of the Lower Ceratite Limestone, without any further precision.
Fig. 11. Biostratigraphical subdivisions of the Griesbachian, Dienerian and earliest Smithian of the Salt Range and correlation with zonations of other regions. Thick vertical black bars indicate uncertainty intervals for correlations. The O. sakuntala beds, indicated with question marks and a dashed frame, correspond to a horizon identified only in the condensed horizon of Nammal Nala (LMH-3, Fig. 4). See text for details (RZ: Regional Zone).
Fig. 12. Biostratigraphical correlations of Chiddru, Wargal, Amb and Nammal Nala sections. Note the diachronism of the Lower Ceratite Limestone between the different sections.
Fig. 13. Range chart showing the biostratigraphical distribution of Griesbachian and Dienerian ammonoid species in the Salt Range. Dashed thick lines correspond to species of uncertain age found in the condensed horizon of Nammal Nala (RZ: Regional Zone).
Fig. 14. Range chart showing the biostratigraphical distribution of Griesbachian and Dienerian ammonoid genera (grouped by families) in the Salt Range. Dashed thick lines correspond to genera of uncertain age occurring in the condensed horizon of Nammal Nala. Grey thick lines correspond to virtual occurrences (RZ: Regional Zone).
Fig. 15. Ghazalaites roohii n. gen. et n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (D = conch diameter, H = whorl height, W = whorl width, U = umbilical diameter).
Fig. 16. Boxplots for the different species of Gyronites from the Salt Range. Because of allometric growth, these boxplots were calculated excluding specimens smaller than 20 mm in diameter, except for G. frequens which shows a strong allometry and for which specimens smaller than 30 mm in diameter have been excluded. Abbreviations as in Figure 15.
Fig. 17. Gyronites frequens Waagen, 1895. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 18. Gyronites frequens Waagen, 1895. Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 30 mm in diameter have been excluded.
Fig. 19. Gyronites dubius (von Krafft, 1909). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 20. Gyronites plicosus Waagen, 1895. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 21. Gyronites plicosus Waagen, 1895. Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 20 mm in diameter have been excluded from this analysis.
Fig. 22. Boxplots for the different species of Ambites from the Salt Range. Because of allometric growth, these boxplots were calculated excluding specimens smaller than 25 mm in diameter. Abbreviations as in Figure 15.
Fig. 23. Ambites discus Waagen, 1895. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Onto-genetic trajectories obtained from sectioned specimen illustrated on Plate 6, figures 17, 18 are shown in black.
Fig. 24. Ambites discus Waagen, 1895. Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 25 mm in diameter have been excluded.
Fig. 25. Ambites atavus (Waagen, 1895). Scatter diagrams of H, W and U, and of H/D, W/D, andU/D (abbreviations as in Fig. 15).
Fig. 26. Ambites atavus (Waagen, 1895). Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 25 mm in diameter have been excluded.
Fig. 27. Ambites radiatus (Brühwiler, Brayard, Bucher & Guodun, 2008). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 28. Ambites radiatus (Brühwiler, Brayard, Bucher & Guodun, 2008). Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 25 mm in diameter have been excluded.
Fig. 29. Ambites bojeseni n.sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 30. Ambites subradiatus n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 31. Ambites superior (Waagen, 1895). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 10, figures 32, 33 are shown in black.
Fig. 32. Ambites superior (Waagen, 1895). Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 25 mm in diameter have been excluded.
Fig. 33. Ambites lilangensis (von Krafft, 1909). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 34. Ambites bjerageri n.sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 35. Boxplots for the different species of Paranoritidae from the Dienerian of the Salt Range. Because of allometric growth, these were calculated only for specimens of 30 to 70 mm in diameter. Abbreviations as in Figure 15.
Fig. 36. Koninckites vetustus Waagen, 1895;. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 14, figures 31–36 are shown in black.
Fig. 37. Koninckites vetustus Waagen, 1895;. Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, specimens smaller than 30 mm in diameter have been excluded.
Fig. 38. Koninckites khoorensis (Waagen, 1895). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 18, figures 10–15 are shown in black.
Fig. 39. Koninckites khoorensis (Waagen, 1895). Comparison between the histograms of H/D, W/D, and U/D for four samples occurring in four successive beds at Nammal Nala. Because of allometric growth, only specimens of 30–70 mm in diameter were taken into account.
Fig. 40. Koninckites khoorensis (Waagen, 1895). Histograms of H/D, W/D, and U/D (abbreviations as in Fig. 15). Because of allometric growth, only specimens of 30–70 mm in diameter were taken into account.
Fig. 41. Radioceras truncatum (Spath, 1934). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15), with comparison of the Dienerian of the Salt Range with faunas from the Smithian of the Salt Range (data from Brühwiler et al. 2012) and from the Dienerian and Smithian of Mud (Spiti Valley, India; data from Brühwiler et al. 2010a).
Fig. 42. Pashtunites kraffti (Spath, 1934) n. gen. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 43. Awanites awani n. gen. et sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 44. Koiloceras sahibi n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 45. Boxplots for the different species of Proptychitidae from the Dienerian of the Salt Range. Because of allometric growth, these boxplots were calculated without the specimens smaller than 30 mm in diameter. Abbreviations as in Figure 15.
Fig. 46. Bukkenites sakesarensis n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 47. Proptychites lawrencianus (de Koninck, 1863) sensu Waagen, 1895;. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 22, figures 14, 15 are shown in black.
Fig. 48. Proptychites ammonoides Waagen, 1895;. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 49. Boxplots for the different species of Mullericeratidae and Hedenstroemiidae from the Dienerian of the Salt Range. Abbreviations as in Figure 15.
Fig. 50. Mullericeras spitiense (von Krafft, 1909). Scatter diagrams of H and W, and of H/D and W/D (abbreviations as in Fig. 15).
Fig. 51. Mullericeras shigetai n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 52. Mullericeras niazii n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 53. Ussuridiscus varaha (Diener, 1895). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 54. Ussuridiscus ensanus (Krafft, 1909 in Krafft & Diener). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 55. Kingites davidsonianus (de Koninck, 1863). Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 31, figures 3, 4 are shown in black.
Fig. 56. Kingites korni Brühwiler, Ware, Bucher, Krystyn & Goudemand, 2010a. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15).
Fig. 57. Clypites typicus Waagen, 1895;. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). The two values for the umbilicus width were measured on internal moulds, the umbilicus being otherwise closed by an umbilical lid.
Fig. 58. Pseudosageceras simplelobatum n. sp. Scatter diagrams of H, W and U, and of H/D, W/D, and U/D (abbreviations as in Fig. 15). Ontogenetic trajectories obtained from sectioned specimen illustrated on Plate 33, figures 4, 5 are shown in black. In addition to the ontogenetic trajectory of the sectioned specimen, a single additional value for the umbilicus width was measured on an internal mould, the umbilicus being otherwise occluded.
Fig. 59. Pseudosageceras simplelobatum n. sp. Histograms of H/D and W/D (abbreviations as in Fig. 15).

Chapter 2

Fig. 1. A, Palaeogeographical map of the Early Triassic with the palaeoposition of Spiti (modified after Brayard et al. 2006). B, Map of India showing the location of Spiti. C, Location map of the three studied areas (open stars) in Spiti District (modified after Brühwiler et al. 2012).
Fig. 2. A, General view of the exposures near Mud, with location of the two studied sections. B, Close-up view of the Permian-Triassic boundary and of the Lower Limestone Member of Mud Top Section. C, Photo of Mud Bottom Section (section M05 of Krystyn et al. 2007a). Note the thickness difference of the Lower Limestone Member compared with the Top Section.
Fig. 3. Section Mud Bottom: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 4. Composite section Mud Top: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 5. Section Guling Village: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 6. Photo of the section Guling River. Note the décollement horizon, with a thick layer of irregular limestone beds replaced laterally by shales.
Fig. 7. Section Guling River: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 8. Section Tilling: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 9. Photo of the section Lalung Cliff. Note the abundance of limestone beds compared with the other sections. The Griesbachian–Dienerian Boundary is here not well defined as no Griesbachian or earliest Dienerian faunas were found. Hence, it is based on the lithological change only.
Fig. 10. Section Lalung Cliff: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 11. Section Lalung Ridge 1: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 12. Section Lalung Ridge 2: lithostratigraphy, ammonoid occurrences and biostratigraphy.
Fig. 13. Biostratigraphic subdivisions of the Dienerian and earliest Smithian of Spiti and correlation with the previously established zonation for this area and the zonation established for the Salt Range (RZ: regional zone).
Fig. 14. Biostratigraphic correlation of the different studied sections, ordered from South to North.
Fig. 15. Range chart showing the biostratigraphical distribution of Dienerian ammonoid species in Spiti (RZ: regional zone).
Fig. 16. Range chart showing the biostratigraphical distribution of Dienerian ammonoid genera (grouped by families) in Spiti (RZ: regional zone).
Fig. 17. Ghazalaites roohii Ware & Bucher 2018b. Scatter diagrams of H/D, W/D, and U/D (D = conch diameter, H = whorl height, W = whorl width, U = umbilical diameter).
Fig. 18. Gyronites frequens Waagen, 1895. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 19. Ambites discus Waagen, 1895. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 20. Ambites bojeseni Ware & Bucher, 2018b. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 21. Ambites nyingmai n. sp. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 22. Ambites lilangensis (von Krafft 1909). Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 23. Ambites lilangensis (von Krafft, 1909). Cumulative histograms of H/D, W/D, and U/D (abbreviations as in Fig. 17). Because of growth allometries, specimens of <25 mm in diameters have been excluded from this analysis. The label ‘normal’ on each histogram indicates that a Lilliefors test performed on the total population (Spiti plus Salt Range) does not allow rejecting the null hypothesis of normality at a confidence interval of 95%.
Fig. 24. Vavilovites meridialis n. sp. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 25. Koninckites vetustus Waagen, 1895. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 26. Koninckites khoorensis (Waagen, 1895). Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 27. Proptychites ammonoides Waagen, 1895. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 28. Mullericeras niazii Ware & Bucher, 2018b. Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 29. Ussuridiscus ensanus (von Krafft, 1909). Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 30. Kingites davidsonianus (de Koninck, 1863). Scatter diagrams of H/D, W/D, and U/D (abbreviations as in Fig. 17).
Fig. 31. Pseudosageceras simplelobatum Ware & Bucher, 2018b. Scatter diagrams of H/D and U/D (abbreviations as in Fig. 17).