High-resolution biochronology and diversity dynamics of the Early Triassic
ammonoid recovery: The Smithian faunas of the Northern Indian Margin
Thomas Brühwiler
a,
⁎, Hugo Bucher
a,
⁎, Arnaud Brayard
b
, Nicolas Goudemand
a
a
Paläontologisches Institut und Museum, Universität Zürich, Karl Schmid-Strasse 4, 8006 Zürich, Switzerland
b
UMR CNRS 5561 Biogéosciences, Université de Bourgogne, 6 Bd. Gabriel, 21000 Dijon, France
abstract article info
Article history:
Received 22 February 2010
Received in revised form 1 September 2010
Accepted 3 September 2010
Available online 9 September 2010
Keywords:
Early Triassic recovery
Smithian
Ammonoids
Northern Indian Margin
Biochronology
Diversity
Based on new collections of abundant and well preserved material from the Salt Range (Pakistan), Spiti
(Northern India) and Tulong (South Tibet), several recent studies focused on the taxonomic revision and
detailed biostratigraphy of Smithian ammonoids. In this work, biochronological data for these three well-
documented basins are analyzed by means of the Unitary Associations method, resulting in a biochronological
scheme of unprecedented high-resolution for the Smithian of the Northern Indian Margin (NIM). Data for
each basin are first processed separately, thus yielding three local biochronological zonations. Then, the three
sequences are processed together as a regional three-section data set for the construction of an inter-basin
sequence at the NIM level. The latter zonation comprises 16 Unitary Associations grouped into 13 zones for
the entire Smithian. Analysis of ammonoid diversity dynamics based on this new highly resolved time frame
highlights (i) a marked diversification during the early Smithian, (ii) a severe extinction during the late
Smithian, and (iii) an overall very high turnover throughout the Smithian. At a global spatial scale and stage
resolution, the diversity of Smithian ammonoid genera appears surprisingly high, as highlighted by a previous
study. It is shown that at a smaller geographic scale and with the most highly resolved time frame, Smithian
ammonoids of the NIM reached their explosive diversity peak essentially through extremely high turnover
rates rather than through a classic diversification process of high origination rates coupled with low extinction
rates. Based on recently published U/Pb ages, regional apparent total rates of origination and extinction of
more than 100 species per My can be inferred for the Smithian ammonoids of the NIM.
© 2010 Elsevier B.V. All rights reserved.
1. Introduction
In the aftermath of the end-Permian mass extinction, ammonoids
were among the fastest clades to recover. The recent analysis of a
global diversity data set of ammonoid genera from the Late
Carboniferous to the Late Triassic shows that Triassic ammonoid
genera actually reached levels of diversity higher than in the Permian
less than 2 million years after the Permian–Triassic Boundary (PTB)
(Brayard et al., 2009). The evolution of Early Triassic ammonoids was
neither a smooth, nor a gradual process. It is characterized by the
following main features: (i) a very low diversity in the Griesbachian
(early Induan), (ii) a moderate diversity increase in the Dienerian
(late Induan), (iii) an explosive radiation in the early Smithian (early
Olenekian), (iv) a late Smithian extinction event followed by (v) a
second explosive radiation during the early Spathian (late Olenekian,
Brayard et al., 2006), and (vi) another significant diversity drop
around the Spathian/Anisian boundary (Bucher, 1989; Brayard et al.,
2009). This generic diversification scenario is coeval with a marked
biogeographical structuring and strong latitudinalisation of faunas
(Brayard et al., 2007).
Several localities on the Northern Indian Margin (NIM) such as the
Salt Range (Pakistan), Kashmir, Spiti (Northern India) and Tulong
(South Tibet) have long been known for their abundant Early Triassic
ammonoid faunas (Mojsisovics et al., 1895; Waagen, 1895; Diener,
1897; Frech, 1905; Krafft and Diener, 1909; Diener, 1913; Spath, 1934;
Schindewolf, 1953; Kummel, 1966, 1970; Wang and He, 1976; Guex,
1978; Bando, 1981). Among these, the Salt Range and Spiti have
played central roles in the development of the Early Triassic time
scale. However, the majority of Smithian ammonoids from the NIM
areas were hitherto insufficiently known due to small numbers of
individuals and/or poor preservation, all of which influencing the
definition of taxa and as a consequence, their biostratigraphic and
biogeographic distributions. Moreover, the exact stratigraphic occur-
rences of many taxa remained either approximate or unknown. Based
on new collections of abundant and well preserved material, a
number of recent studies focused on the taxonomic revision and
detailed biostratigraphy of Smithian ammonoids from the Salt Range
(Brühwiler et al., in press-b), from Spiti (Krystyn et al., 2007a,b;
Brühwiler et al., 2010b, in press-a) and from Tulong (Brühwiler et al.,
2010a)(Fig. 1). These studies have resulted in the most complete and
Palaeogeography, Palaeoclimatology, Palaeoecology 297 (2010) 491–501
⁎ Corresponding authors. Tel.: +41 44 634 26 98; fax: +41 44 634 49 23.
E-mail addresses: bruehwiler@gmx.net (T. Brühwiler), hugo.fr.bucher@pim.uzh.ch
(H. Bucher).
0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2010.09.001
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