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465
Chapter 17
Permian-Triassic Extinctions
and Rediversifications
Arnaud Brayard and Hugo Bucher
© Springer Science+Business Media Dordrecht 2015
C. Klug et al. (eds.), Ammonoid Paleobiology: From macroevolution to paleogeography,
Topics in Geobiology 44, DOI 10.1007/978-94-017-9633-0_17
A. Brayard ()
UMR CNRS 6282 Biogéosciences, Université de Bourgogne, 6 boulevard Gabriel,
21000 Dijon, France
e-mail: arnaud.brayard@u-bourgogne.fr
H. Bucher
Paläontologisches Institut und Museum, Universität Zürich, Karl-Schmid Strasse 4,
8006 Zürich, Switzerland
e-mail: hugo.fr.bucher@pim.uzh.ch
17.1 Introduction
At the boundary between the Paleozoic and Mesozoic eras (~ 252 myr), the end-
Permian mass extinction was the most devastating global-scale event ever recorded,
resulting in the loss of more than 90 % of marine species (Raup 1979) and the dis-
appearance or severe reduction in diversity of typical Paleozoic organisms (e.g.,
trilobites, tabulate and rugose corals, brachiopods). The ecological recovery of
the benthos is traditionally assumed to have spanned the entire Early Triassic (i.e.
~ 5 myr), thus strikingly contrasting with that of pelagic environments and their
dwellers. Whether or not this difference is the result of a selective preservation bias
against the benthos cannot be excluded. However, extreme diversity fluctuations of
nekto-pelagic organisms (e.g., ammonoids and conodonts) during the entire Early
Triassic indicate major environmental upheavals in the ocean in the wake of the
end-Permian extinction(s). In support of markedly unstable Early Triassic times,
several major events are known from the sedimentary, geochemical and palynologi-
cal records (e.g., Payne et al. 2004, 2010; Galfetti et al. 2007a, b, c; Hermann et al.
2011, 2012; Sun et al. 2012; Grasby et al. 2013; Romano et al. 2013; Fig. 17.1a),
suggesting profound global changes in climate, sea-level and oceanic geochemistry
(e.g. anoxia, euxinia, acidification). The initial low resolution time frames of these
recurrent environmental deterioration events after the Permian-Triassic boundary
(PTB) crisis were therefore first lumped into a “delayed recovery” model which
is still the standard in effect in some recent reviews (e.g., Chen and Benton 2012).
466 A. Brayard and H. Bucher
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46717 Permian-Triassic Extinctions and Rediversifications
Contrasting with this early view, recent analyses of nekto-pelagic taxa such as am-
monoids and conodonts document an explosive Early Triassic rediversification (Or-
chard 2007; Brayard et al. 2009c). Indeed, although ammonoids were among the
organisms most affected by the PTB mass extinction, Triassic ammonoids actually
reached levels of diversity much higher than in the Permian less than ~ 1.5 myr after
the PT boundary (Brayard et al. 2009c). In this chapter, we provide a brief overview
of the present state of our knowledge of the ammonoid record around the PTB crisis
and during the recovery interval.
17.2 Late Permian Events
Four major clades of Permian ammonoids (Goniatitida, Prolecanitida, Tornocera-
tida and Ceratitida; Fig. 17.1a; e.g., Ruzhencev 1960; Glenister and Furnish 1981;
Leonova 2002, 2011) are globally characterized by relatively slow and uncoupled
origination and extinction dynamics (Brayard et al. 2009c). Goniatitida were the
dominant group during the Early-Middle Permian interval (e.g., Ruzhencev 1960;
Glenister and Furnish 1981; Leonova 2002, 2011). Following a diversity peak dur-
ing the Middle Permian, all ammonoid groups show a protracted, two-step decline
in diversity during the Late Permian (Capitanian and Changhsingian extinctions;
Glenister and Furnish 1981; Stanley and Yang 1994; Zhou et al. 1996). The first ex-
tinction event in the Capitanian is now generally agreed to have been triggered by the
eruption of the Emeishan flood basalts (Bond et al. 2010). Remarkably, Ceratitida
did not follow this trend and flourished during the Wuchiapingian with the notable
rapid diversification of the morphologically-singular Otocerataceae (Brayard et al.
2009c; Leonova 2002, 2011; Zakharov and Abnavi 2013). This time interval also
corresponds to a marked restriction of global ammonoid morphological disparity
(e.g., Saunders et al. 2008; Villier and Korn 2004; Leonova 2005). With blossoming
Xenodiscidae and Pseudotirolitidae, Ceratitida also largely embodied the taxonomic
diversity of Changhsingian ammonoids. The PTB mass extinction was most probably
triggered by eruption of the Siberian traps (Reichow et al. 2009; Svensen et al. 2009).
Fig. 17.1 a Total ammonoid generic richness ( black bold line: all ammonoids; color lines: major
ammonoid groups; Permian bold line: data from Goniat.org; Permian dotted line: alternate data
from the Ammon database (Korn and Ilg 2007); Triassic bold line modified after Brayard et al.
(2009c) based on an updated database) and mean Chao2 estimate of the overall generic richness
with its 95 % Confidence Interval (large circles with vertical bars; see Brayard et al. 2015). PTB:
Permian-Triassic boundary. E.T.: Early Triassic. Note that the end-Smithian ammonoid extinction
event discussed in the text is not illustrated here due to its short time duration. b Chronostrati-
graphic subdivisions of the Early Triassic (radiometric ages by Ovtcharova et al. 2006; Galfetti
et al. 2007b and Shen et al. 2011) with simplified trends of geochemical (δ13Ccarb; data from
Galfetti et al. 2007b) and Tethyan relative temperature fluctuations during this period (data from
Romano et al. 2013 [black line] and Sun et al. 2012 [grey line]). c Temporal distribution of some
Early Triassic representative families (modified after Tozer 1981; Brayard et al. 2006, 2009c).
Ammonoid illustrations from Brayard and Bucher (2008), Brayard et al. (2013) and Guex et al.
(2010). d Simplified trend of the formation of a latitudinal diversity gradient during this period
(modified after Brayard et al. 2006, 2007b, 2009b)
468 A. Brayard and H. Bucher
The Goniatitida and Tornoceratida completely disappeared with this event (Glenis-
ter and Furnish 1981; Tozer 1981; Leonova 2002, 2011) and apparently, only one
Prolecanitida genus ( Episageceras) survived the crisis (Kummel 1972; Tozer 1981;
Dagys and Ermakova 1996; Leonova 2011; Zakharov and Abnavi 2013). Among the
Ceratitida, only two main ammonoid clades survived across the PT boundary: the
Otoceratidae and Xenodiscidae. Otoceras rapidly disappeared after the mass extinc-
tion, before the end of the Griesbachian. However, after a gap spanning the entire
Dienerian, Otocerataceae probably had their final appearance with Proharpoceras
in the Smithian (Brayard et al. 2007a). Thus, with very exceptions ( Episageceras,
Otoceras, Proharpoceras) and a still ambiguous case (Sagecerataceae) (e.g., Glenis-
ter and Furnish 1981; Tozer 1981; Becker and Kullman 1981; Brayard et al. 2007a;
McGowan and Smith 2007), Triassic ammonoids are usually agreed to root into a
single and morphologically very simple clade, the Xenodiscidae and are therefore
interpreted as a monophyletic clade (Tozer 1981; Brayard et al. 2006; McGowan and
Smith 2007). Following this hypothesis, all Mesozoic ammonoids are consequently
derived from the xenodiscids, which went unscathed across the PTB mass extinction.
17.3 Early Triassic Events
Early Triassic times are commonly divided into four stages or substages of highly
uneven duration (Griesbachian, Dienerian, Smithian and Spathian), which are very
well defined by ammonoid zones and events (Fig. 17.1b; see Jenks et al. 2015 and
Monnet et al. 2015). Based on ammonoids, the PTB is traditionally defined by the
first occurrence of Otoceras (e.g., Tozer 1994, 2003; Dagys and Ermakova 1996;
Zakharov 2002; Shevyrev 2006). Although knowledge of Griesbachian and Die-
nerian ammonoids is still limited, proptychitids and meekoceratids originated dur-
ing the Dienerian. In their first steps, these Dienerian originations did not lead to
any spectacular taxonomic diversification (e.g., Tozer 1974; Shevyrev 2001; Brüh-
wiler et al. 2008; Ware et al. 2011; Fig. 17.1c). The beginning of the Smithian is
defined by the origination of new families such as the highly speciose Flemingitidae
and Kashmiritidae. Ammonoid richness first peaked during the middle Smithian
through the evolution of extremely short-lived species (Brayard et al. 2009c; Brüh-
wiler et al. 2010). The beginning of the Spathian witnessed an explosive radiation
rooted in the latest Smithian Xenoceltites. Tirolitidae first quickly diverged from
Xenoceltites and formed the basal group of from which the next radiation that began
in the early Spathian and continued onward at a very high pace.
The tempo of recovery after the PTB mass extinction has recently been estimated
thanks to new U-Pb radiometric ages allowing for more accurate and precise time
calibrations (Ovtcharova et al. 2006; Galfetti et al. 2007b; Shen et al. 2011). The
rediversification was explosive for some marine groups such as the ammonoids
(Brayard et al. 2009c) and conodonts (Orchard 2007), spanning less than ~ 1.5 myr
(Fig. 6.21a). Ammonoids reached levels of taxonomic richness in the Smithian
that were already much higher than those of the Permian (see also Tozer 1981;
Leonova 2002). Their recovery did not follow Sepkoski’s (1978) classical logistic
46917 Permian-Triassic Extinctions and Rediversifications
diversification model, for which major assumptions are required about the carrying
capacity of the environment, but it instead has been re-interpreted as a hierarchical
model periodically interrupted by brief but significant extinction events (e.g., end-
Smithian; Brayard et al. 2009c).
The global first-order trend in increasing ammonoid diversity was accompanied
by a progressive change from cosmopolitan to latitudinally-restricted distribution
during the Early Triassic with the formation of a clear latitudinal diversity gra-
dient during most of the Smithian and Spathian sub-stages (Fig. 17.1d; Brayard
et al. 2006, 2015). Marked intertropical faunal exchanges across the Tethys and
Panthalassa are obvious at that time with the occurrence of identical ammonoids on
opposite sides of Panthalassa, illustrating latitudinally-restricted faunal exchanges
during the Smithian (Brayard et al. 2007b, 2009a, b, 2013; Jenks et al. 2010), and
the Spathian (Galfetti et al. 2007b; Guex et al. 2010; Monnet et al. 2013).
The global recovery trend was not a continuous process. It was interrupted at
least once during a brief episode of ammonoid cosmopolitanism combined with a
marked extinction event during the end-Smithian (e.g., Tozer 1982; Dagys 1988;
Brayard et al. 2006). This extinction event was the most important one within the
entire Triassic and its intensity compares with that of the PTB extinction. Only four
species-poor families (the xenoceltitids and the sageceratids, but also virtually the
palaeophyllitids and the proptychitids) survived the late Smithian extinction.
Similarly, conodonts reached their highest Triassic generic diversity during the
middle Smithian, but only a very few species went through the end-Smithian crisis.
On land, floras underwent a drastic ecological turnover with a middle Smithian
spore peak comparable to the end-Permian one, followed by an early gymnosperm
recovery during the late Smithian (Hermann et al. 2011). An abrupt, global change
from hygrophytic to xerophytic associations characterizes the Smithian-Spathian
boundary (Galfetti et al. 2007c; Hermann et al. 2011). The global carbon isotope
record also reached a marked negative peak during the middle Smithian, followed
by an abrupt positive shift in the late Smithian (e.g., Payne et al. 2004; Galfetti et al.
2007b). In the Tethys, the oxygen isotope record from biogenic phosphate tends to
track the carbon isotope record and indicates a temperature drop of ca. 7.5 °C close
to the Smithian-Spathian boundary (Romano et al. 2013).
It is well known by ammonoid workers (e.g., Kummel and Steele 1962) that
the Early Triassic ammonoid radiation is represented by numerous homeomorphic
taxa. Trends in morphological disparity and richness were especially decoupled
during the Griesbachian and Dienerian with persisting low disparity values in the
Dienerian whereas richness shows a weak increase (McGowan 2004, 2005; Brosse
et al. 2013). The first disparity peak occurred early in the Smithian (Brosse et al.
2013). The end-Smithian extinction had obvious consequences with a marked con-
traction of the previously occupied morphospace. The Spathian corresponds to a
second disparity peak with a morphospace analogous to the early-middle Smithian.
However, Spathian superfamilies occupied more restricted portions of the morpho-
space (Brosse et al. 2013).
Ammonoid recovery during the Early Triassic therefore appears as the com-
bined outcome of (i) the classical rapid refilling of a vacated ecospace after the
mass extinction, and (ii) the successive extinction events and recurrent stressful
470 A. Brayard and H. Bucher
environmental conditions that may have enhanced their high turnover rates. Most
likely explanations for the end-Smithian extinction call upon the combined con-
sequences of the concentration of carbon dioxide of volcanic origin (e.g., Galfetti
et al. 2007b) and sea-level changes. Whatever the precise cause(s), it had a deep im-
pact on the biotic rediversification, especially for ammonoids, conodonts and plant
ecological assemblages as well. We cannot exclude the possibility that a few other
Early Triassic biotic events are still hidden due to the insufficient knowledge of
intervals such as the Griesbachian and Dienerian. However, significant results and
new questions on the Early Triassic recovery will certainly arise from further stud-
ies of ammonoids and their spectacular evolutionary rebounds, which are hardly
reconcilable with the alleged persistence of globally devastated ecosystems.
Acknowledgments We thank D. Korn, J. Jenks and K. Bylund for their constructive comments
and suggestions. The CNRS INSU Interrvie supported A.B. for his study. This is also a contribu-
tion to the ANR project AFTER (ANR-13-JS06-0001). H.B. acknowledges the support of the
Swiss National Science Foundation (project 200021_135446).
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