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Proc. R. Soc. B (2008) 275, 759–765

doi:10.1098/rspb.2007.1370
Published online 15 January 2008

Recovery from the most profound

mass extinction of all time
Sarda Sahney* and Michael J. Benton
Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queen’s Road, Bristol BS8 1RJ, UK
The end-Permian mass extinction, 251 million years (Myr) ago, was the most devastating ecological event
of all time, and it was exacerbated by two earlier events at the beginning and end of the Guadalupian, 270
and 260 Myr ago. Ecosystems were destroyed worldwide, communities were restructured and organisms
were left struggling to recover. Disaster taxa, such as Lystrosaurus, insinuated themselves into almost every
corner of the sparsely populated landscape in the earliest Triassic, and a quick taxonomic recovery
apparently occurred on a global scale. However, close study of ecosystem evolution shows that true
ecological recovery was slower. After the end-Guadalupian event, faunas began rebuilding complex trophic
structures and refilling guilds, but were hit again by the end-Permian event. Taxonomic diversity at the
alpha (community) level did not recover to pre-extinction levels; it reached only a low plateau after each
pulse and continued low into the Late Triassic. Our data showed that though there was an initial rise in
cosmopolitanism after the extinction pulses, large drops subsequently occurred and, counter-intuitively, a
surprisingly low level of cosmopolitanism was sustained through the Early and Middle Triassic.
Keywords: alpha; diversity; mass extinction; Permian; Triassic; recovery

1. INTRODUCTION the Kungurian and Roadian stages, 271 million years

The end-Permian mass extinction was the most dramatic (Myr) ago. This marks the turnover between Early
event to impact life on Earth (Erwin 1990, 2006; Benton Permian pelycosaur-rich faunas to those dominated by
2003; Benton & Twitchett 2003). On land and sea, life was therapsids and pareiasaurs and has been termed ‘Olson’s
nearly extinguished, ecosystems were devastated and gap’ (Lucas & Heckert 2001) or ‘Olson’s extinction’
many long-lived lineages disappeared. Most studies (Sahney & Benton in preparation), depending on whether
hitherto have focused on the effects of this extinction on it is seen as a failure of fossil preservation or a real
marine life. It has been more difficult to determine the ecological event. Frustratingly, tetrapod footprint faunas
effects on land because rock successions are typically less of this time show an even longer hiatus than the skeletal
complete, and only two major sedimentary basins have so evidence, so that the validity of each interpretation is
far yielded extensive faunas spanning the boundary: the difficult to verify (Lucas 2004).
Karoo Basin of South Africa and the South Urals Basin The impact of the end-Permian event was devastating.
of Russia. In the sea, the level of species loss was 80–96%, and
During the last two decades, our understanding of the blastoid echinoderms, tabulate and rugose corals, grapto-
end-Permian extinction has increased significantly. It is lites, trilobites, eurypterids, acanthodians and placoderms
now widely believed that this devastating event was caused disappeared entirely (Hallam & Wignall 1997). On land,
by large-scale volcanism in Siberia, which led to major the dominant Glossopteris flora was replaced, eight orders
atmospheric changes and the collapse of ecosystems of insects became extinct (Labandeira & Sepkoski 1993)
worldwide (Benton 2003). Earlier evidence suggested and two-thirds of tetrapod families were lost (Benton
that this extinction was prolonged through the Late 1989). The only tetrapod lineages to survive were
Permian (Erwin 1990; Teichert 1990), but hints of an procolophonoids, dicynodonts, and presumably theroce-
earlier event, noted first among terrestrial vertebrate phalians, cynodonts, and archosauromorphs, and their
faunas (Benton 1985, 1989; King 1991), have been Triassic recovery was slow (Benton et al. 2004).
confirmed (Retallack et al. 2006) as the same end- The return of diversity after an extinction event can be
Guadalupian extinction of marine organisms, already viewed in terms of taxonomy ( Niklas et al. 1983;
noted independently ( Jin et al. 1994; Stanley & Yang Labandeira & Sepkoski 1993; Benton 1995), ecology
1994; Rohde & Muller 2005). During both extinction
(Bambach 1985; Bambach et al. 2007) or morphology
events, diversity losses were not simply a reflection of
( Foote 1995; Roy & Foote 1997). In this study, we are
increased rates of extinction but also of depressed
concerned with the recovery of taxonomic diversity in the
origination rates (Benton 1998; Bambach et al. 2004).
context of ecological diversity.
The terrestrial vertebrate record of the Permian is also
There are several meanings for the term ‘recovery’ after
confounded by an additional earlier event, or gap, between
mass extinctions. Past studies have revealed that faunal
revival after a devastating ecological event may follow a
* Author for correspondence (s.sahney@bristol.ac.uk). pattern similar to ecological succession (Sole et al. 2002),
Electronic supplementary material is available at http://dx.doi.org/10. and recovery may be considered as the point at which
1098/rspb.2007.1370 or via http://journals.royalsociety.org. the model is complete and the new ecosystem is stable.

Received 4 October 2007 759 This journal is q 2008 The Royal Society
Accepted 12 December 2007
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760 S. Sahney & M. J. Benton Recovery from mass extinction

45 2. MATERIAL AND METHODS

40 Diversity is most commonly assessed by tallying the number
no. of tetrapod families

of taxa on a global scale (Benton 1985, 1995, 1998; Padian &

35
Clemens 1985); such studies provide a synoptic overview but
30 2
are subject to the vagaries of sampling. This is especially true
25 in the fossil record where patterns of global diversity often
20 reflect patterns of fossiliferous rock availability (Raup 1972;
15 Peters & Foote 2001; Smith 2001; Smith & McGowan 2005).
10 1 3 In this study, we use an alternative approach by sampling
5 individual fossil communities (Alroy et al. 2001). By
examining well-preserved and well-studied faunas, the
0
280 270 260 250 240 230 taxonomic and ecological recovery of communities after the
Ar K R W Ca Wu Ch I O An L Cr Permo-Triassic extinction event can be examined more
accurately, and the problems of geological bias between
Figure 1. Global diversity (dashed line) and mean alpha
diversity (solid line) of Permo-Triassic tetrapod families. time bins are largely avoided.
Extinctions are labelled as 1, Olson’s extinction; 2, end- Data were collected for 69 tetrapod communities from the
Guadalupian extinction; and 3, end-Permian extinction. Artinskian to the end of the Carnian and stored in a relational
Geological stages (Gradstein & Ogg 2004) are as follows: Ar, database management system, the alpha diversity database
Artinskian; K, Kungurian; R, Roadian; W, Wordian; Ca, (ADD). We surveyed all fossil tetrapod findings from the
Capitanian; Wu, Wuchiapingian; Ch, Changhsingian; I, Permian and Triassic and chose only those localities/forma-
Induan; O, Olenekian; An, Anisian; L, Ladinian; Cr, Carnian. tions with a high quality of preservation, a completeness of
community and a thorough collection and publication
On both the scale of modern ecological recovery and history. The study is therefore global and we made no
recovery from mass extinction, disaster (‘weedy’ or geographical restrictions. Inevitably, however, we could not
generalist) taxa are known to insinuate themselves find suitably substantial faunas from every time bin from
into empty guilds, pushing the boundaries of their every continent: the Early Permian sample is dominated by
geographical range and ecospace. Early Triassic terrestrial North American localities, the Middle and Late Permian and
ecosystems are clearly dominated by a small number Early Triassic time bins by sections in Russia (Pechora Basin,
of genera, most notably the dicynodont Lystrosaurus, Moscow Basin, South Urals Basin) and southern Africa
which accounted for approximately 90% of terrestrial (Karoo Basin, Ruhuhu Valley), with additional sites from
vertebrates (Benton 1983). Disaster taxa then experienced Greenland, Australia, Madagascar, Antarctica and Germany.
rapid turnover in the time immediately following the Sampling in the Middle and Late Triassic is broader, with
event, later giving way to more specialized organisms localities from Western Europe, Russia, South America,
(Benton 2003). North America, Africa, Madagascar, India and China.
Recovery can also be viewed as a return to pre- The time scale employed was that of Gradstein & Ogg
extinction conditions, for example the numerical recovery (2004) and abbreviations for time units were taken from
of taxa. Globally, this type of recovery happened quickly Benton (1993). Sedimentary environment, climate and
after the end-Permian event by the Olenekian biome were refined based on the discussion with H. Falcon-
(250–245 Myr ago), but then the diversity fell again Lang (2006, personal communication).
(figure 1), either as a result of displaced disaster taxa that The classification of orders and families was taken from
had filled empty guilds or the devastation caused by Benton (1993, 2005). Ambiguous taxonomic information
another extinction pulse at the end of the Olenekian. In was included in the database but was tagged, so that the
contrast, the ecological recovery of tetrapods and plants analyses could be run with and without the unclear data. The
was slow, and lost guilds and trophic levels were not structure allowed for the inclusion of taxa identified as nomina
readily refilled (Retallack et al. 1996; Benton et al. 2004; dubia as well as specimens labelled ?, aff., cf. or otherwise
Grauvogel-Stamm & Ash 2005). indeterminate or unidentified. Stratigraphical ranges and
The third interpretation defines recovery as the point geographical distributions of families were taken from Benton
when an ecological equilibrium is reached, even if this (1993) and updated using Carroll et al. (1998), and Schoch &
balance is unlike that of the pre-extinction fauna. Milner (2000). Ecomorphs based on size and diets were taken
Though faunal turnovers occurred continually before from Benton (1996).
and after the end-Permian event, it is difficult to say
when the ecosystem reached stability, i.e. a time of high
biodiversity, low turnover, resistance to invaders and a 3. EXTINCTION AND RECOVERY IN THE MIDDLE
complex trophic structure (Sole et al. 2002). Immedi- TO LATE PERMIAN
ately after the end-Permian event, amphibians made a Three distinct extinction pulses were responsible for the
relatively quick recovery, specifically the families Capi- mass extinction of tetrapods in the Permian and Triassic:
tosauridae and Trematosauridae, which filled the role Olson’s extinction; the end-Guadalupian event; and the
of semi-aquatic predators. Among reptiles, therapsids end-Permian event. Olson’s extinction, in the Early
filled disaster taxa roles but their dominance gave way Guadalupian (Roadian, Wordian), reveals an extended
to archosauromorphs. period of low diversity when worldwide two-thirds of
In this paper, we present the first community- terrestrial vertebrate life was lost. Global diversity rose
scale analysis of tetrapod recovery from the end- dramatically in the Capitanian, probably the result of
Permian extinction and the preceding Guadalupian disaster taxa filling empty guilds, only to fall again when
extinction events. the end-Guadalupian event caused a diversity drop in the

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Recovery from mass extinction S. Sahney & M. J. Benton 761

diversification of tetrapod families

0

–1

–2

–3

–4

–5
320 300 280 260 240
Miss Penn Cisur Guad Lopin E.Tri M.Tri L.Tri

Figure 2. Rate of alpha diversification (the first derivative of diversity through time) of mean number of tetrapod families in
communities, calculated stage-by-stage from the Mississippian to the Late Triassic; error bars cannot be applied. Miss,
Mississippian; Penn, Pennsylvanian; Cisur, Cisuralian; Guad, Guadalupian; Lopin, Lopingian; E. Tri, Early Triassic; M. Tri,
Middle Triassic; L. Tri, Late Triassic.

Wuchiapingian. Globally, terrestrial vertebrates recovered small, 60% medium and 20% large tetrapods in the Late
the high familial diversity of the Artinskian (39 non- Permian follows (figure 3a).
singleton families found worldwide) in the Changhsin- Olson’s extinction was a dramatic extinction ‘trough’
gian, the last stage of the Permian (figure 1), only to be that is a prolonged period of very low diversity after a long
devastated again by the end-Permian event. and sustained diversity rise and probably the result of
The communities in this study are globally dispersed prolonged environmental stress. Both Olson’s extinction
and despite the differences in latitude and climate, they are and the end-Guadalupian extinction experience a dramatic
fairly well constrained in their diversity. Community change in community diet proportions, body size and
diversity falls dramatically during Olson’s extinction and distinct faunal turnovers, from ‘pelycosaur’-rich commu-
diverse Artinskian faunas, which featured large amniote nities to those dominated by the basal therapsid dinoce-
predators and herbivores such as Dimetrodon and Diadectes, phalians and finally giving way to more derived therapsids
and a variety of semi-aquatic tetrapods, including Eryops, and pareiasaurs. Global and ecological diversities recover to
Archeria and Ophiacodon, disappear. The mean number of pre-extinction levels by the end of the Permian. However,
families in Early Guadalupian communities is reduced to a alpha diversity does not recover by reaching a pre-extinction
paltry 13% of the Artinskian high. A detailed look at the balance; rather a new equilibrium is reached, significantly
Russian Permian sequence confirms that the diversity drop lower than the diversity of the Artinskian.
was severe; only two families crossed the boundary from
the Wordian to the Capitanian, but recovery began
immediately with the origination of five families in the 4. ACROSS THE PERMO-TRIASSIC BOUNDARY:
latter stage (Benton et al. 2004). Globally, alpha diversity ECOSYSTEM RESTRUCTURING AND RECOVERY
does not fully recover; it slowly reaches a plateau (approx. (a) A fast global recovery and a slow
57% of Artinskian diversity) by the end of the Permian community recovery
(figure 1). The pattern of diversification is similar at all The global diversity rose sharply after each extinction
taxonomic levels and taxonomic proportions are consistent pulse, probably the result of disaster taxa filling empty
across all time periods. guilds. After the end-Permian event, this rapid refilling
The ecological impact of the Guadalupian events is resulted in a return to pre-extinction taxonomic diversity
catastrophic; 8 (out of a possible 12) guilds are lost from by the Olenekian. However, this did not last, as there was a
the Artinskian high of 10 guilds. These are recovered in subsequent loss of nine families (figure 1).
the last stages of the Permian before being devastated Until the Carnian, community diversity never reached
again by the end-Permian event (figure 3c). A dramatic the highest observed in Artinskian faunas. Though the
change in diet type also occurs: proportions of piscivores, end-Permian extinction pulse had the most dramatic
insectivores, predators and browsers are thrown out of global effect, it does not appear to have impacted the
balance during each extinction pulse (figure 3b). During diversity within individual communities as badly as the
the last two stages of the Permian, there is a movement first Guadalupian pulse. It seems to have ‘thinned out’
towards pre-extinction diet proportions, but again the families rather than destroyed them completely and the
end-Permian event disrupts this recovery. Body size is average number of tetrapod families lost is only 1.5 from
affected in a similar manner: the proportion of small, each community (figure 1).
medium and large animals is thrown out of balance and A look at the community diversification rate helps
recovery towards a pre-extinction balance of roughly 20% explain the slow recovery of tetrapods in the Triassic

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762 S. Sahney & M. J. Benton Recovery from mass extinction

(a) an average of one family was added every 4 Myr. Then for
100 four stages after Olson’s extinction, this rate increased to
body size proportion of community

one family added every 3 Myr. But oddly, for the four
80 stages immediately following the end-Permian extinction,
a period of 35 Myr, the familial diversification rate dips in
60 and out of the negative realm and results in an average of
one additional family every 25 Myr. Outside the three
40 mass extinction events, this is the lowest rate of tetrapod
diversification since their origin in the Frasnian.
20
(b) Ecological recovery
0 Communities were slow to recover ecologically as well
280 270 260 250 240 230
Ar K R W Ca Wu Ch I O An L Cr as numerically, and body size and diet ratios show
disruptions at all extinction pulses (figure 3). The Permian
extinctions created a ‘Lilliput effect’ (Urbanek 1993) in
(b) 100 which few large species survived and, in addition, many
small species were also extinguished. The large animals
diet proportion of community

80 included the disaster taxon Lystrosaurus that dominated

terrestrial communities, capitosaurs and Proterosuchus,
60 which were populous in semi-aquatic environments.
Further into the Early Triassic, larger animals became
40 more common as dicynodonts such as Kannemeyeria
replaced Lystrosaurus, and large carnivores reappeared
20 including Cynognathus, Garjainia and Erythrosuchus.
The end-Permian event dramatically restructured
0 communities with the loss of browsers and predators and
280 270 260 250 240 230
an increase in piscivores. The loss of browsers is, no doubt,
Ar K R W Ca Wu Ch I O An L Cr
linked to changes in vegetation. A rapid loss of sediment-
binding vegetation was responsible for a basin-wide
change from low-energy meandering streams to high-
(c) 12
energy braided rivers in the Early Triassic of Russia, South
10 Africa, Australia, India and Spain ( Newell et al. 1999;
Ward et al. 2000; Benton in press). Also, the 7 Myr ‘coal
no. of occupied guilds

8 gap’ is the result of an insufficient amount of plant material

to form coal deposits, and hence little food for large
6
browsing animals. Permian levels of plant diversity were
4 not reached again until the Late Triassic (230 Myr ago;
Retallack et al. 1996). Tracking diet proportions through
2 the first five stages of the Triassic shows a clear trend
towards reaching a pre-extinction balance of communities
0 250 dominated by predators and browsers with a smaller
280 270 260 240 230
Ar K R W Ca Wu Ch I O An L Cr proportion of piscivores and insectivores (figure 3b).
Overall, the recovery of the eight guilds lost in the
Figure 3. Ecological diversity of Permo-Triassic tetrapod
Guadalupian occurred in 10–15 Myr, relatively quickly
communities. (a) Size (light green, large; light yellow,
medium; sky blue, small) and (b) diet (pink, browsers; light compared with the original 100 Myr it took to colonize
green, predators; light yellow, insectivores; sky blue, pisci- them by the first tetrapods (Sahney & Benton in
vores) are expressed as a percentage of the total community. preparation), perhaps because sufficient representatives
(c) The guilds are defined by body size and inferred diet. of the relevant lineages and guilds survived the event in
Error bars cannot be applied as the number of occupied some form. These broad guilds had just been refilled by
guilds is not a mean number, rather a cumulative count of the the end of the Permian, when the end-Permian extinction
guilds filled in each stage. Size of the animals is defined as pulse caused the loss of two major guilds (small piscivores
small (with snout-vent length (SVL) less than 150 mm), and small browsers). These two are not regained by the
medium (SVL from 150 mm to 1.5 m) and large (SVL end of the Carnian (table 1).
greater than 1.5 m). Geological stages are the same as given in
A slow Triassic recovery may initially seem at odds with
figure legend 1.
Botha & Smith’s (2006) analysis of the Karoo Basin,
which indicates an initial rapid recovery of the tetrapod
(figure 2). Since the Viséan, there have been three distinct fauna after the end-Permian event. They noted that nearly
periods of diversification interrupted by three major half (43%) of the Early Triassic genera appeared ‘rapidly’,
extinctions, in the Late Pennsylvanian, Middle Permian that is within 20 m of the Permo-Triassic boundary. As
and End Permian. During the periods of diversification they state, however, they are documenting the onset of
(with the exception of the Moscovian), fewer than one recovery, rather than the establishment of mature
family was added every Myr, representing a slow but ecosystems. Botha & Smith (2006) defined recovery as a
steady rate of community-level diversification. For the ‘new balance’ rather than a return to pre-existing
four stages following the Late Pennsylvanian extinction, conditions, something that could not be observed in the

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Recovery from mass extinction S. Sahney & M. J. Benton 763

1.0

cosmopolitanism of tetrapod families

0.8
3

0.6 1

2
0.4

0.2

0
320 300 280 260 240
Miss Penn Cisur Guad Lopin E.Tri M.Tri L.Tri

Figure 4. Cosmopolitanism of tetrapods through the Carboniferous, Permian and Triassic. Cosmopolitanism (C ) is measured as
mean alpha diversity ðT Þ divided by global diversity (Tt ), according to the formula CZ T =Tt . Note the overall decline of
cosmopolitanism through this time interval, perhaps related to increasing taxonomic and ecological diversity of tetrapods, but
also note the coupled rises and falls in cosmopolitanism following major extinction events, especially (1) Olson’s extinction,
(2) the end-Guadalupian extinction and (3) the end-Permian extinction. Abbreviations are the same as given in figure legend 2.

Table 1. Guilds (defined by body size and diet) filled by Permo-Triassic tetrapods.

piscivores insectivores browsers predators

midpoint guilds guild
stage (Myr ago) S M L S M L S M L S M L occupied loss/gain

Artinskian 280 Y Y Y Y Y Y Y Y Y Y 10
Kungurian 273.1 Y Y Y Y Y Y Y 7 K3
Roadian 269.3 Y Y Y 3 K4
Wordian 266.9 Y Y 2 K1
Capitanian 263.1 Y Y Y Y Y Y 6 4
Wuchiapingian 257.1 Y Y Y Y Y Y Y Y Y 9 3
Changhsingian 252.4 Y Y Y Y Y Y Y Y Y Y 10 1
Induan 250.35 Y Y Y Y Y Y Y 7 K3
Olenekian 247.35 Y Y Y Y Y Y Y Y 8 1
Anisian 241 Y Y Y Y Y Y Y Y Y 9 1
Ladinian 232.5 Y Y Y Y Y Y Y Y 8 K1
Carnian 222.25 Y Y Y Y Y Y Y Y 8 0

Karoo alone as this sequence begins in the Guadalupian, ecological models, which predict that when endemic/
after the first extinction pulse. regional faunas are devastated, the survivors, especially
A longer sequence of tetrapod faunas is found in the disaster taxa, become cosmopolitan, at least in the
Southern Urals of Russia, where a continuous record of immediate aftermath of a mass extinction (Hallam &
tetrapod faunas spanning 30 Myr from the Kungurian to Wignall 1997; Benton 2003; Erwin 2006). This is well
the Ladinian reveals low taxonomic and ecological illustrated among tetrapods by Lystrosaurus, a bulky
diversity in the Early Triassic. Even after 15 Myr of herbivore that thrived in the harsh arid conditions of the
rebuilding ecosystems, many guilds were still unfilled, Induan. Scroungers such as Lystrosaurus were then rapidly
such as those of small piscivores, small insectivores, large overtaken by the evolution of more stable regionally
herbivores and top predators (Benton et al. 2004). distinct faunas that were to establish the longer lasting
ecosystems of the later Triassic.
(c) Cosmopolitanism There is a repeated pattern of coupled rises and falls in
Cosmopolitanism is the degree to which a species or a cosmopolitanism during times of extinction, though in
clade is distributed worldwide; it can be measured simply each case the fall is far more dramatic than the rise,
as the mean alpha diversity of communities divided by creating an overall downward trend. Cosmopolitanism
global diversity. The Pennsylvanian extinction, Guadalu- was slow to recover in the Triassic, rising just 7% from the
pian pulses and end-Permian event experience significant Olenekian to the Carnian. There is no other time in
rises in cosmopolitanism followed by dramatic losses of tetrapod history that cosmopolitanism is sustained at such
40–50% (figure 4). This rise and fall of cosmopolitanism a low level for so long. Continued low cosmopolitanism in
following an extinction event is in keeping with established the Triassic is difficult to envisage owing to the existence of

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764 S. Sahney & M. J. Benton Recovery from mass extinction

the global supercontinent, Pangaea and equable climates: Benton, M. J. 1998 The quality of the fossil record of
terrestrial organisms should have been able to migrate vertebrates. In The adequacy of the fossil record (eds S. K.
nearly worldwide without major barriers. Donovan & C. R. C. Paul), pp. 269–303. New York, NY:
Wiley.
Benton, M. J. 2003 When life nearly died: the greatest mass
extinction of all time. London, UK: Thames & Hudson.
5. CONCLUSIONS Benton, M. J. 2005 Vertebrate palaeontology. Oxford, UK:
The Permo-Triassic extinction events halted the growth of Blackwell Science.
tetrapod communities. Extinction in the Guadalupian had Benton, M. J. In press. The end-Permian mass extinction:
a much more severe impact at the community level than events on land in Russia. Proc. Geol. Assoc. 119.
the terminal end-Permian event. Faunas were recovering Benton, M. J. & Twitchett, R. J. 2003 How to kill (almost) all
from the Guadalupian events at reasonably high rates and life: the end-Permian extinction event. Trends Ecol. Evol.
ecologically communities had recovered when the end- 18, 358–365. (doi:10.1016/S0169-5347(03)00093-4)
Permian event hit. Though globally tetrapods recovered Benton, M. J., Benton, M. J., Tverdokhlebov, V. P. & Surkov,
quickly, the dramatic restructuring that occurred at the M. V. 2004 Ecosystem remodelling among vertebrates at
community level was not permanent and communities did the Permian–Triassic boundary in Russia. Nature 432,
not recover numerically or ecologically in the Early and 97–100. (doi:10.1038/nature02950)
Middle Triassic. It would not be until the great diversity of Botha, J. & Smith, R. M. H. 2006 Rapid vertebrate
the Late Triassic, which included dinosaurs, pterosaurs, recuperation in the Karoo Basin of South Africa following
the end-Permian extinction. J. Afr. Earth Sci. 45, 502–514.
crocodilians, rauisuchids, aetosaurs, rhynchosaurs, trilo-
(doi:10.1016/j.jafrearsci.2006.04.006)
phosaurs, sphenodonts, amphibians and mammals, some
Carroll, R. L., Bossy, K. A., Milner, A. C., Andrews, S. M. &
30 Myr after the end-Permian event, that terrestrial Wellstead, C. F. 1998 Encyclopedia of paleoherpetology:
tetrapod community diversity was restored. Lepospondyli. München, Germany: Dr Friedrich Pfeil.
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advice of Saswati Bandyopadhyay, Max Langer, Graeme 110190.000441)
Lloyd, Rainer Schoch, and Anne Warren, on Triassic Erwin, D. H. 2006 Extinction: how life on Earth nearly ended
communities. This work was partially funded by NERC 250 million years ago. Princeton, NJ: Princeton University
grant NE/C518973/1 to M.J.B. and supported by grants to Press.
S.S. from the Bob Savage Fund, the Natural Environment Foote, M. 1995 Morphological diversification of Paleozoic
Research Council, the Palaeontological Association, and the crinoids. Paleobiology 21, 273–299.
University of Bristol Alumni Foundation. Gradstein, F. M. & Ogg, J. G. 2004 Geologic time scale
2004—why, how, and where next! Lethaia 37, 175–181.
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