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2007 The Paleontological Society. All rights reserved. 0094-8373/07/3304-0004/$1.

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Paleobiology, 33(4), 2007, pp. 547573
Macroecological responses of terrestrial vegetation to climatic
and atmospheric change across the Triassic/Jurassic boundary in
East Greenland
Jennifer C. McElwain, Mihai E. Popa, Stephen P. Hesselbo, Matthew Haworth,
and Finn Surlyk
Abstract.The magnitude and pace of terrestrial plant extinction and macroecological change as-
sociated with the Triassic/Jurassic (Tr/J) mass extinction boundary have not been quantied using
paleoecological data. However, tracking the diversity and ecology of primary producers provides
an ideal surrogate with which to explore patterns of ecosystem stability, collapse, and recovery and
to explicitly test for gradual versus catastrophic causal mechanisms of extinction.
We present an analysis of the vegetation dynamics in the Jameson Land Basin, East Greenland,
spanning the Tr/J extinction event, from a census collected paleoecological data set of 4303 fossil
leaf specimens, in an attempt to better constrain our understanding of the causes and consequences
of the fourth greatest extinction event in earth history. Our analyses reveal (1) regional turnover
of ecological dominants between Triassic and Jurassic plant communities, (2) marked structural
changes in the vegetation as reected by potential loss of a mid-canopy habit, and (3) decline in
generic-level richness and evenness and change in ecological composition prior to the Tr/J bound-
ary; all of these ndings argue against a single catastrophic causal mechanism, such as a meteorite
impact for Tr/J extinctions. We identify various key ecological and biological traits that increased
extinction risk at the Tr/J boundary and corroborate predictions of meta-population theory or plant
ecophysiological models. These include ecological rarity, complex reproductive biology, and large
leaf size.
Recovery in terms of generic-level richness was quite rapid following Tr/J extinctions; however,
species-level turnover in earliest Jurassic plant communities remained an order of magnitude high-
er than observed for the Triassic. We hypothesize, on the basis of evidence for geographically ex-
tensive macrofossil and palynological turnover across the entire Jameson Land Basin, that the na-
ture and magnitude of paleoecological changes recorded in this study reect wider vegetation
change across the whole region. How exactly these changes in dominance patterns of plant primary
production affected the entire ecosystem remains an important avenue of future research.
Jennifer C. McElwain.* Department of Geology, The Field Museum, 1400 South Lake Shore Drive, Chicago,
Illinois 60605. E-mail: Jennifer.McElwain@ucd.ie
Mihai E. Popa. University of Bucharest, Faculty of Geology and Geophysics, Department of Geology and
Palaeontology, 1, Nicolae Balcescu Avenue, 010041, Bucharest. E-mail: mihai@mepopa.com
Stephen P. Hesselbo. Department of Earth Sciences, University of Oxford, Oxford, OX1 3PR, United King-
dom. E-mail: Stephen.Hesselbo@earth.ox.ac.uk
Matthew Haworth. UCD School of Biology and Environmental Science, University College Dublin, Na-
tional University of Ireland, Beleld, Dublin 4, Ireland. E-mail: Matthew.Haworth@ucd.ie
Finn Surlyk. Institute of Geography and Geology, University of Copenhagen, ster Voldgade 5-7, DK-1350
Copenhagen, Denmark. E-mail: FINNS@geol.ku.dk
*Present address: UCD School of Biology and Environmental Science, University College Dublin, National
University of Ireland, Beleld, Dublin 4, Ireland
Accepted: 13 June 2007
Introduction
The nature, causes, and consequences of the
Triassic/Jurassic (Tr/J) mass extinction event
(200 Myr ago) have received increasing atten-
tion over the past decade. Sepkoskis (1981)
original global compendium of marine faunal
extinction rates classied the Tr/J boundary
as one of the big ve extinction events in
earth history. Estimates of the magnitude of
diversity loss across the boundary vary from
group to group and are dependent on the
scale of study. Stage-level compilations for
marine families and genera (Sepkoski 1981,
1993) indicate losses of 23% and 50% respec-
tively. At the regional or single locality level,
extinction magnitudes of species are extreme-
ly high: 80% of terrestrial plant species in
Greenland and Sweden (Harris 1937; Mc-
Elwain et al. 1999) and 42% of terrestrial ver-
tebrates families in North America (Olsen et
548 JENNIFER C. MCELWAIN ET AL.
al. 1987), as well as widespread extinction of
ammonites (Newell 1963; Tozer 1979), bi-
valves (McRoberts and Newton 1995), radio-
larians (Tipper et al. 1994; Carter and Hori
2005), and coral reefs (Kiessling 2001, 2005).
In contrast, other authors have questioned
whether the boundary can be characterized
unequivocally as a mass extinction. Hallam
(2002) has argued that the tempo of extinction
was gradual rather than catastrophically rap-
id, whereas others (Tanner et al. 2004) suggest
that most of the apparent biodiversity losses
across the Tr/J boundary are due to biases or
artifacts of sampling or poor stratigraphic
control. Reanalysis of Sepkoskis global ma-
rine faunal data has also demonstrated that
low origination rates were more responsible
than high extinction rates for Tr/J biodiversity
loss (Bambach et al. 2004).
Uncertainties regarding the nature and tem-
po of the Tr/J boundary extinctions are fur-
ther complicated by the fact that most biotic
records spanning the RhaetianHettangian
interval are based on presence-absence data,
which can articially indicate a catastrophic
event if taxonomic groups are oversplit or
bias interpretation in favor of a gradual event
owing to taphonomic control of last occur-
rences (Signor III and Lipps 1982). Presence-
absence data sets also limit our understanding
of the ecological and physiological selectivity
of extinctions. For instance, is there evidence
for gradual ecosystem decline or instability
prior to the extinction event at the Tr/J
boundary? Does the extinction event repre-
sent a biotic threshold response to long-term
gradual forcing or was it truly catastrophic?
Were the taxa that suffered extinction ecolog-
ically important dominants within ecosys-
tems or rare? In the absence of paleoecological
data these critical questions about the re-
sponses of terrestrial plant communities
across the Tr/J boundary remain unan-
swered. Without paleoecological data it is also
difcult to decipher potential forcing mecha-
nisms of biotic turnover, many of which make
explicit predictions about the nature and tem-
po of macroecological response. To address
these uncertainties we have undertaken a de-
tailed paleoecological study of terrestrial
plant communities through Rhaetian and Het-
tangian strata of the Kap Stewart Group in Ja-
meson Land, East Greenland. Froma database
of 4304 census-collected macrofossil plant
specimens we have investigated the tempo of
macroecological change across the Tr/J
boundary and tracked stability, collapse, and
recovery of plant communities, the primary
production and therefore foundation of terres-
trial ecosystems in this region. We dene cen-
sus-collected as follows: where every fossil
specimen discovered within a specic fossil
plant bed was collected within a standardized
time frame for all plant beds.
We have conducted this paleoecological
analysis in the context of a dramatically
changing global environment throughout the
TriassicJurassic interval, with peak environ-
mental changes coinciding with the Tr/J
boundary, as reected by a negative carbon
isotopic excursion in both organic and inor-
ganic carbon (Palfy et al. 2001; Ward et al.
2001; Hesselbo et al. 2002; Guex et al. 2004;
Galli et al. 2005). We have used relative abun-
dance data and two measures of biodiversity
(evenness and richness) collected from nine
fossil plant beds to assess whether there was
a gradual decline in ecosystem functioning
prior to the Tr/J boundary, or whether an ob-
served 80% species-level extinction at the
boundary (Harris 1937; McElwain et al. 1999)
represents a geologically instantaneous event.
We have also investigated compositional
changes (including the degree of heterogene-
ity) in terrestrial vegetation and the ecological
and physiological selectivity of the Tr/J event
in an attempt to understand better the causal
mechanisms of Tr/J boundary oral turnover.
Abiotic Context for Tr/J Extinctions
Causal mechanisms for Tr/J extinctions re-
main unresolved. Suggested forcing factors
for Tr/J biodiversity loss include climatic and
environmental disturbance due to a cata-
strophic meteorite impact (CAMP) (Olsen et
al. 2002), gradual climatic and environmental
change associated with emplacement of the
Central Atlantic Magmatic Province (Marzoli
et al. 1999, 2004; McElwain et al. 1999; Hes-
selbo et al. 2002), catastrophic release of meth-
ane due to methane hydrate destabilization
resulting in global warming (Palfy et al. 2001;
549 TRIASSIC/JURASSIC VEGETATION DYNAMICS
FIGURE 1. Stable carbon isotopic (
13
C) record from fossil wood (Hesselbo et al. 2002) (A), cuticle (McElwain et al.
1999) (B), and a paleoatmospheric carbon dioxide record (McElwain et al. 1999) (C), plotted against height of the
section at Astarteklft (cf. Fig. 3) across the Triassic (Rhaetian)/Jurassic (Hettangian) boundary. The carbon isotopic
data in A are all derived from fossil wood collected at Astarteklft. The cuticle material in B is derived from As-
tarteklft and other sections throughout the Jameson Land Basin (see McElwain et al. 1999 for details).
Beerling and Berner 2002), and sea-level
change involving a fall followed rapidly by a
rise (Hallam 1997). Global cooling has also
been invoked as a potential extinction mech-
anism (Hubbard and Boulter 2000) although
supporting evidence for such a mechanism is
open to serious challenge.
Irrespective of the exact causal mechanism,
stable carbon isotopic records have identied
a pronounced double negative excursion of
23 coincident with the boundary in East
Greenland (McElwain et al. 1999; Hesselbo et
al. 2002), Hungary (Palfy et al. 2001), England
(Hesselbo et al. 2002), the United States (Guex
et al. 2004; Ward et al. 2006), Italy (Galli et al.
2005, 2006), and Canada (Williford et al. 2006).
These records clearly document a major per-
turbation of the carbon cycle, which was glob-
al in nature, and they suggest major environ-
mental upheaval in the latest Rhaetian with
maximum environmental change coinciding
with the Tr/J boundary. Hesselbo et al. (2002)
interpreted the single carbon isotope excur-
sion apparent in the Jameson Land succession
as an amalgamation of an initial and a
main isotope excursion observed in ex-
panded marine sections.
In this paper we explore in detail the ter-
restrial plant communities in the region of As-
tarteklft, East Greenland, in the context of
the stable carbon isotopic composition from
fossil wood in the same section (McElwain et
al. 1999; Hesselbo et al. 2002) (Fig. 1A,B). Al-
though it is difcult to infer directly from iso-
topic records how the climate or atmospheric
composition was changing we interpret the
isotopic prole to provide an integrated re-
cord of environmental upheaval across the in-
terval. We therefore interpret any signicant
global deviation in
13
C (2 from average
background levels) as a perturbation in the
carbon cycle associated with a change in en-
vironmental conditions. We also investigate
vegetation dynamics in the context of a likely
fourfold increase in paleoatmospheric CO
2
across the Tr/J boundary from levels that
were approximately three times higher than
present before the boundary, derived using
the stomatal proxy approach (McElwain et al.
1999), and a 16C regional climatic warming,
550 JENNIFER C. MCELWAIN ET AL.
FIGURE 2. Map of the Jameson Land Basin showing lo-
cations of the fossil plant localities and position of pa-
leolake.
inferred from a coupled ocean atmosphere
general circulation model forced with this CO
2
reconstruction (Huynh and Poulsen 2005).
However, an estimated regional warming of
16C is remarkably high and likely to be over-
estimated, as their model does not take into
account the possible effect of higher SO
2
levels
due to CAMP volcanism (which would cool
climates) on global or regional climate warm-
ing. Furthermore, stomatal density-based CO
2
records are required from additional genera
to Ginkgoales, following recent procedures
for other time intervals (i.e., McElwain et al.
2005) to test the delity of the existing Ginkgo-
based record. Paleo-CO
2
records based on
multiple independently calibrated genera
with temporally overlapping stratigraphic
ranges are more robust, as they minimize cal-
ibration uncertainty that can be introduced
due to genetic variability in stomatal frequen-
cy within and between genera, such as that
demonstrated by (Cantor et al. 2006).
Material
Kap Stewart Flora. In a seminal series of
monographic papers published between 1926
and 1937 TomHarris documented a Rhaetian
Hettangian aged fossil ora of over 200 spe-
cies from approximately 13 localities in Ja-
meson Land, East Greenland (Fig. 2). This fos-
sil ora is represented by a rich assemblage of
bryophytes, pteridophytes and gymnosperms
(Harris 1926, 1931, 1932a,b, 1935, 1937). Two
distinct plant macrofossil biozones are rec-
ognized within the Kap Stewart Group: a
Rhaetian assemblage zone characterized by
the presence of Lepidopteris ottonis and a Het-
tangianSinemurian assemblage zone with
Thaumatopteris brauniana (Harris 1937). A
80% species-level turnover of plant macro-
fossils occurs between the highest occurrences
of L. ottonis zone taxa and the lowest occur-
rences of T. brauniana zone taxa (Harris 1937).
This oral transitional zone was used by
Harris to dene the Tr/J boundary. Subse-
quent eldwork (20002004) has shown that
the rst appearance of Thaumatopteris brauni-
ana zone elements occur contemporaneously
with the last appearances of Lepidopteris zone
taxa in Bed 5 of our current study locality, As-
tarteklft. This level coincides with the most
negative carbon isotopic values in the stable
carbon isotopic record from the same section
(Fig. 3) (Hesselbo et al. 2002). Palynological
studies of the Kap Stewart Group have de-
ned two microoral zones that parallel the
macrooral biozones of Harris and correlate
with established Rhaetian aged Rhaetipollis-
Limbosporites zone and Hettangian aged Pi-
nuspollenites-Trachysporites zones of Europe,
Canada, and Svalbard (Pedersen and Lund
1980). For these reasons, and in the absence of
a global boundary stratotype section and
point (GSSP) for the Tr/J boundary, we inter-
pret the top of Bed 5 as representing the
Tr/J boundary at Astarteklft.
Geological and Depositional Setting. The
RhaetianHettangian aged Kap Stewart
Group in Jameson Land was deposited in en-
vironments that ranged from uvial to lacus-
trine (Dam and Surlyk 1993b) (Table 1). The
group comprises three formations (Surlyk
2003): the predominantly conglomeratic and
sandy Innakajik Formation at the base, depos-
ited in coarse-grained alluvial plain environ-
ments; a mixed sandy and shaley Primulaelv
Formation in the middle, deposited in a delta-
plain setting; and a mixed sandy and shaley
Rhaetelv Formation at the top, deposited in a
551 TRIASSIC/JURASSIC VEGETATION DYNAMICS
lacustrine setting. The plant fossils collected
for this study come exclusively from the Pri-
mulaelv Formation at Astarteklft (Figs. 2, 3).
The succession at Astarteklft comprises
mainly sandstone (7580%), of ne to coarse
grade and commonly texturally and compo-
sitionally immature (Dam and Surlyk 1993b).
Shaley units, which constitute 2025%of the
succession, are made up of pale blocky mud-
stones ranging in thickness up to a maximum
of 4 m, and dark-gray leaf-rich shale in beds
1 m. Poorly developed black coal and coa-
ly mudstone occurs about the middle of the
formation. Rootlets are present at the base of
the coal, and at several other horizons in the
mudstone facies. Thin beds of siderite occur in
the more organic-rich shaley layers. Most of
the mudstone units are traceable across the ex-
posure at individual localities and evidently
have lateral extents on the order of tens to
hundreds of meters. Mudstone beds also oc-
cur within channelized sandstone bodies.
Thin sand laminae and wave-ripple structures
have not been observed in the mudstone facies
at Astarteklft.
The thick sandstones were laid down from
river channels that were commonly several
meters deep (Sykes 1974; Dam and Surlyk
1993a,b). The textural and mineralogical im-
maturity of the sand indicates a local source
terrain that was undergoing strong physical
erosion; in the case of the Hurry Inlet (east Ja-
meson Land) localities this was likely to have
been the Caledonian basement that is current-
ly exposed in Liverpool Land to the east (Fig.
2). Neither discrete erosional channel margins
nor mud-plugged abandoned channels are
commonly observed, indicating rapid and
continuous lateral migration of channels dur-
ing deposition. The thin coarsening-upward
sandstone sequences that form wings to ad-
jacent channel lls represent sheet splay or
crevasse splay deposits formed during river
ood episodes (Damand Surlyk 1993b). These
are the primary facies in which plant fossils
occur at Astarteklft (sheet splay in Table 1, SS
in Fig. 3). A secondary setting for the plant
fossils is in sediments deposited at upward
transitions from channel sand facies into
oodplain facies (abandoned channel in Table
1, AC in Fig. 3). These are interpreted to have
been shallow pools developed in semi-aban-
doned sand-lled channels.
The predominant gray color of the mud-
stone units, combined with moderate lateral
extent and sporadically rooted nature, indi-
cates deposition on a permanently or semi-
permanently waterlogged oodplain or in an
interdistributary lacustrine bay. The differ-
ence between oodplain and interdistributary
lacustrine bay environments is one of degree
of connection to open lake conditions. In our
study, in the Hurry Inlet area, interpretation of
these facies as oodplain is preferred because
sedimentary structures indicative of open wa-
ters (e.g., wave ripples) are absent, palynofa-
cies are exclusively of terrestrial origin (Kop-
pelhus 1997), and there is no observed inter-
digitation of alluvial or delta plain with la-
custrine facies. The true coal (coal swamp
[CS]), a third category of plant bed in Jameson
Land Basin, possibly only occurs at one strati-
graphic level (equivalent with the level of Bed
6 at Astarteklft; Table 1, Fig. 3) but is not ful-
ly developed at Astarteklft. Only rarely did
the oodplain areas dry out enough for oxi-
dizing conditions to become established, as
shown by the volumetrically subordinate pur-
ple-gray mudstone. The abundance of green-
clay cementation of the tops of sandstone beds
indicates the common development of anaer-
obic early diagenetic conditions beneath the
waterlogged oodplain.
The oras preserved in these three plant-
bed settings are likely to be of somewhat dif-
ferent origins. Floodplain deposits and coal
swamps are likely to represent mainly autoch-
thonous assemblages, containing mostly in
situ representatives of the oodplain and
swamp communities, respectively (Gastaldo
1989). Sheet or crevasse splays contain autoch-
thonous assemblages from local wetland and
oodplain communities, combined with
allochthonous plant material derived from
channel levees and possibly from plant com-
munities further upstream (Gastaldo 1989;
Spicer 1989). These types of deposits are
thought to provide some of the most accurate
representations of the overall regional source
vegetation despite taphonomic biases that
tend to overestimate upper delta or alluvial
plain communities and underrepresent those
552 JENNIFER C. MCELWAIN ET AL.
553 TRIASSIC/JURASSIC VEGETATION DYNAMICS
TABLE 1. Fossil plant collection statistics from 2002 eld season.
Bed Age
Depositional
environment
No. rock slabs
collected
No. fossil
leaves recorded
No.
reproductive
specimens
recorded
Bed 8 Jurassic (Hettangian) Abandoned channel 150 221 0
Bed 7 Jurassic (Hettangian) Abandoned channel 170 864 10
Bed 6 Jurassic (Hettangian) Coal swamp 80 128 1
Bed 5 Tr/J boundary Sheet splay 332 1023 (320)* 3 (2)*
Bed 4 Triassic (Rhaetian) Sheet splay 412 876 103
Bed 3 Triassic (Rhaetian) Sheet splay 127 525 0
Bed 2 Triassic (Rhaetian) Sheet splay 157 258 4
Bed 1.5 Triassic (Rhaetian) Sheet splay 43 62 0
Bed 1 Triassic (Rhaetian) Sheet splay 87 224 1
* Numbers in brackets refer to number of specimens collected census style from Bed 5. The higher number in Bed 5 refers to the total number of
specimens collected using census techniques, collected directly from the bed and from fallen blocks directly under the bed.

FIGURE 3. Measured section from Astarteklft, East Greenland from Hesselbo et al. (2002), updated based on fur-
ther eldwork in 2002 and 2004. Biostratigraphy and lithostratigraphy from Harris (1937), Pedersen and Lund
(1980), Dam and Surlyk (1993a,b) and Surlyk (2003). The fossil plant beds located in the present study are shown
in boldface; Beds 0, 5.5, and 9 and the moss bed have not yet been investigated in detail. Harriss (1937) plant
bed names are also shown, together with his barometer-based estimated distance below the Jamesoni Horizon (Har-
ris scale). We re-located Harriss beds on the basis of position in the section and on the contained macrooras.
Interpreted depositional environments are indicated in italics. SS, sheet splay; CS, coal swamp; AC, abandoned
channel.
of the local oodplain (Ferguson 1985; Gas-
taldo 1989; Spicer 1989). The abandoned chan-
nels, which according to sedimentological
analysis were formed by avulsion, would be
expected to preserve predominantly parauto-
chthonous plant communities growing in
close proximity to the channel, with only very
rare occurrences of allochthonous compo-
nents transported from upstream (Behrens-
meyer et al. 2000). Fossil plant assemblages in
these deposits would therefore most likely
provide insights only into plant communities
in the immediate region of Astarteklft.
Taphonomic Considerations
Taphonomic studies have shown that un-
transported and undecayed litter from tem-
perate and relatively low diversity subtropical
oodplain forests provides an accurate indi-
cation of both the richness and the dominance-
diversity relationship of the source forest
(Burnham 1989; Burnham et al. 1992). It is
noteworthy that the majority of these live-
dead studies have been carried out on angio-
sperm dominated plant communities, with
only one detailed study to date on a forest
where a gymnosperm was co-dominant (Tax-
odium-Acer swamp [Burnham et al. 1992]).
Nonetheless, observations in different ecolog-
ical settings and hemispheres have revealed
that representation of live species in leaf litter
from both angiosperm- and gymnosperm-
dominated (Taxodium) forests is predominant-
ly controlled by the same factors (e.g., relative
abundance, canopy height, leaf area, distance
from tree) (Burnham et al. 1992; Steart et al.
2005). In the absence of a live-dead study of
broad-leaved conifer dominated vegetation,
we have therefore assumed that the same pri-
mary factors controlled preservation of leaf
litter in the Triassic and Jurassic. General bi-
ases that should be noted include slight over-
representation of taxa with a woody vine (li-
ana) habit (Burnham et al. 1989), and better
representation of larger canopy trees than
smaller sub-canopy taxa (Gastaldo 2001). Bi-
ases due to decay and transport of leaf litter
can also be considerable (Ferguson 1985; Spi-
cer 1989).
Despite these caveats, the macrofossil as-
semblages from Astarteklft are exquisitely
preserved and abundant, primarily autoch-
thonous or parautochthonous and to a lesser
extent allochthonous, and show little evidence
554 JENNIFER C. MCELWAIN ET AL.
for decay or long-distance transport. Further-
more all fossil plant beds were census-collect-
ed to avoid collector bias, and lateral sampling
of each bed was as wide as the nature of ex-
posure would allow, so as to minimize biasing
of the rank-abundance data by very localized
input of leaves from individual plants. Every
bed was therefore laterally sampled across a
minimum of eight to a maximum of 20 me-
ters. Six out of a total of nine fossil plant beds
are isotaphonomic, including all ve of the
Triassic beds and the Tr/J bed (Table 1, Fig.
3). Isotaphonomic assemblages afford the
least-biased means of tracking biodiversity
trends through time (Behrensmeyer et al.
2000; Gastaldo 2001). The large database of
fossil leaf counts from macrofossil assemblag-
es at Astarteklft therefore provides an ideal
study system with which to investigate paleo-
ecological changes in Triassic and Jurassic
vegetation. In the case of the six isotaphon-
omic plant beds, we can explicitly examine
paleoecological changes that occurred leading
up to, and coincident with, the Tr/J extinction
boundary bed, and test gradual versus cata-
strophic mechanisms of oral extinction.
However, because the depositional setting of
the Jurassic plant beds at Astarteklft (two
abandoned channels and one swamp) are dif-
ferent from those of the Triassic (all sheet
splay), we use preliminary fossil plant occur-
rence data from a Jurassic abandoned channel
assemblage at a second Jameson Land eld lo-
cality (South Tancrediaclft [McElwain, I.
Glasspool, Popa, Hesselbo, D. Sunderlin, and
Surlyk unpublished data]) to distinguish the
potential effects of taphonomy from larger-
scale paleoecological patterns across the Tr/J
boundary.
Collection Methods
Macrofossil specimens were collected in
2002 using census style techniques from a to-
tal of nine fossil plant beds at Astarteklft, Ja-
meson Land, East Greenland (Table 1, Fig. 3).
Most these fossil plant beds were originally
discovered by Harris and were relocated by
measuring distance in meters below a pecten-
rich carbonate-cemented sandstone (Jamesoni
Horizon) of the Neill Klinter Group (Fig. 3).
Harris used this as his marker bed throughout
Jameson Land localities to correlate macroo-
ras across the basin (Harris 1937: p. 71). Col-
lection procedures were standardized for all
nine fossil plant beds. Each bed was collected
for a total of 48 hours by excavating four small
quarries, spaced two to ve meters apart lat-
erally, as constrained by the nature of the ex-
posure. Roughly the same amount of bulk
sediment was excavated per bed with the ex-
ception of Beds 1.5 and 6, which were only
census-collected for a total of eight hours each.
All macrofossil specimens excavated during
the collection interval were collected irrespec-
tive of preservation state and are currently
stored in the paleobotanical collections at the
Field Museum, Chicago. Every identiable
macrofossil specimen was identied to genus
in the laboratory; specimens of each genus
were counted and recorded in a fossil leaf oc-
currence matrix (Table 2). We use the term
macrofossil specimen to refer to any iden-
tiable vegetative or reproductive structure
preserved on a rock slab/specimen. Individ-
ual slabs commonly preserved multiple fossil
leaves and reproductive structures from one
or many fossil plant taxa (Table 1). To estimate
the relative abundances of Triassic and Juras-
sic genera at Astarteklft we used a sampling
strategy that represents a hybrid method be-
tween the widely used quadrat method of
counting (Pfefferkorn et al. 1975), which re-
ports the occurrence of a taxon once per quad-
rat irrespective of howmany leaves there were
per quadrat, and those methods that count ev-
ery individual leaf (Wing et al. 1993). Quadrat
methods can inate the relative abundances of
ecologically rare taxa and underestimate the
abundances of dominants (see Wing and
DiMichele 1995 for a full discussion), whereas
individual counting methods are extremely
time consuming and it can be difcult to re-
solve when, and if, leaf fragments should be
counted. We have counted every individual
leaf or leaf fragment per rock slab as one oc-
currence in our abundance matrix (Table 2).
However, in cases where there were more than
ve leaves or ve leaf fragments per rock slab
we recorded a count of ve rather than count-
ing every individual leaf. We recognize that
this counting strategy may underestimate the
relative abundances of the ecological domi-
555 TRIASSIC/JURASSIC VEGETATION DYNAMICS
TABLE 2. Generic abundance (number of occurrences) of macrofossil leaf and reproductive specimens at Astar-
teklft, East Greenland.
Macrofossil taxon Bed 1 Bed 1.5 Bed 2 Bed 3 Bed 4 Bed 5 Bed 6 Bed 7 Bed 8
Anthrophyopsis 0 0 0 0 2 0 0 0 0
Anomozamites 36 20 21 1 237 1 0 4 0
Baiera 14 0 0 7 0 0 0 0 0
Cladophlebis 0 0 9 0 0 3 50 22 12
Clathropteris 0 0 4 0 0 0 0 0 0
Ctenis 0 0 0 0 0 4 0 0 0
Ctenozamites 0 0 0 0 0 0 0 0 0
Cycadocarpidium 0 0 0 0 0 1 0 0 0
Cycadolepis 1 0 4 0 88 1 0 1 0
Czekanowskia 5 0 0 0 0 0 0 442 27
Dictyophyllum 2 19 1 2 144 0 11 0 17
Doratophyllum 0 0 0 41 0 0 0 0 0
Elatocladus 1 4 16 0 1 3 19 0 3
Equisetites 24 0 0 0 10 5 2 0 0
Ginkgoites 14 0 80 0 1 0 0 214 0
Hausmannia 1 0 0 0 0 0 0 0 0
Lepidopteris 1 2 12 0 27 0 0 0 0
Macrotaeniopteris 0 0 0 0 0 5 0 0 0
Marattia 0 0 0 0 0 0 1 0 0
Mattonia 0 0 0 0 0 0 0 0 6
Neocalamites 0 0 0 0 0 11 0 0 0
Nilssonia 7 9 0 3 0 0 0 1 0
Ontheodendron 0 0 0 0 0 3 0 0 0
Pachypteris 0 0 0 0 1 0 0 0 0
Pagiophyllum 0 0 0 0 0 0 1 0 0
Phlebopteris 0 0 0 0 0 0 1 0 34
Podozamites 62 1 28 453 13 519 0 0 44
Pseudoctenis 0 3 0 0 1 28 0 0 0
Pterophyllum 56 3 49 11 440 0 5 0 0
Ptilozamites 0 0 1 0 0 9 1 0 0
Rhinipteris 0 0 35 0 0 0 0 0 2
Sagenopteris 0 0 0 0 0 0 2 0 0
Sphenobaiera 0 0 0 6 0 0 0 181 76
Spiropteris 0 0 0 0 4 0 1 0 0
Stachyotaxus 1 1 2 0 0 416 35 0 0
Taeniopteris 0 0 0 1 1 9 0 0 0
Thaumatopteris 0 0 0 0 0 4 0 0 0
Vardekloeftia 0 0 0 0 0 1 0 0 0
Weltrichia 0 0 0 0 4 0 0 0 0
Wielandiella 0 0 0 0 5 0 0 0 0
nants; however, we feel that this is preferable
to the loss of potentially important ecological
data caused by the ination of abundances of
rare taxa. Furthermore, a sensitivity analysis
comparing our counting strategy for Bed 1 at
Astarteklft with a standard quadrat style
counting approach has revealed only subtle
differences in the resultant relative and rank-
order abundances of fossil leaf genera and in-
signicant differences in standard measures
of biodiversity based on the relative occur-
rence data (Appendix 1). Census collection of
the Kap Stewart Group at Astarteklft result-
ed in a data set of 4303 recorded occurrences
of macrofossil leaf and reproductive speci-
mens, derived from a total of 40 Rhaetian and
Hettangian plant genera.
Paleoecological Analysis
Paleoecological analyses have all been car-
ried out at the genus level. An analysis of Har-
riss original Tr/J macrofossil data (Harris
1937) indicates that 95% of generic occurrenc-
es, from a total of 38 different beds, are mono-
specic. Those genera containing more than
one species occur predominantly in Triassic
(9%) rather than Jurassic plant beds (0.5%)
and are typically represented by two (e.g.,
Ctenis, Equisetites) and rarely three to four spe-
cies (e.g., Pterophyllum) in a single assemblage.
556 JENNIFER C. MCELWAIN ET AL.
From these data we infer that a generic-level
analysis is a fairly robust proxy for analysis at
the species level (which will be the subject of
future work) but that generic-level richness
may underestimate species richness in the Tri-
assic more than in the Jurassic. We note that a
generic-level paleoecological analysis may
also mask interesting species dynamics be-
tween stratigraphic levels. However, these
cannot be resolved until we have a rmer un-
derstanding of the real taxonomic value of leaf
surface micromorphological features (e.g., sto-
matal density and distribution, trichome
abundance, presence and absence) on which
species determinations for the Kap Stewart
Group ora were largely based (Harris, 1926,
1931, 1932a,b, 1937). It is now known than
many of these micromorphological traits are
highly plastic in response to changes in the cli-
matic and atmospheric environment. A num-
ber of species as determined by Harris could
therefore represent a continuum of ecophen-
otypes responding to environmental changes
associated with CAMP volcanic activity, rath-
er than biological species.
Composition. Small changes in the ecolog-
ical composition of plant communities can
greatly alter whole ecosystem processes, bi-
otic interactions, and feedbacks with the cli-
matic system. Signicant functional shifts in
ecosystem processes can result from changes
in the relative dominance of different taxa
without any signicant change in taxonomic
composition. To track changes in generic-level
oral composition and dominance patterns
across the Tr/J boundary, we calculated rela-
tive generic abundances per bed for all 41 gen-
era as a percentage of the total bed abundance
using the occurrence matrix (Table 2).
Detrended correspondence analysis (DCA)
was carried out on the resultant relative abun-
dance matrix (Table 3) using PAST Version
1.33 (Hammer et al. 2001). This eigenvector
technique was developed for detecting oris-
tic gradients in modern botanical data sets
and displaying complex compositional differ-
ences between sites in two dimensions (Hill
and Gauch 1980). Using this technique, sites
or plant beds that are compositionally sim-
ilar, plot close together in ordination space,
whereas those which are compositionally dis-
tinct plot far apart. In the case of this study we
display compositional change using the rst
eigenvector (Axis 1) only. To minimize poten-
tial biases introduced by disarticulated repro-
ductive structures of unknown phylogenetic
afnity, we restricted our investigations of
compositional changes to vegetative macro-
fossils only. DCA was also repeated with the
inclusion and exclusion of genera occurring in
only one bed to test for any distortion effects
that these may introduce in the analyses (Wilf
and Johnson 2004).
Biodiversity. Biodiversity is variously de-
ned by different authors. However, we dene
biodiversity here in terms of both richness (the
number of taxa) and evenness (the equality of
relative abundances among taxa) of the pa-
leovegetation. The more diverse or rich an eco-
system the more likely that it will contain
functional types that are highly productive
(Loreau et al. 2001). Although there are many
exceptions to the rule, richer ecosystems tend
to be more productive as long as climatic and
edaphic variables are favorable. Also the richer
an ecosystem, the more likely it is to with-
stand major abiotic changes, because there is
a larger species pool from which new domi-
nant taxa or new productive functional types
can be recruited.
Evenness is a measure of niche partitioning
and facilitation within an ecosystem (Loreau
et al. 2001). Ecosystems that have many co-
dominant species all contributing to ecosys-
tem function, such as in modern tropical low-
land forests, have a high degree of evenness
and a higher diversity of phenotypic traits.
They are therefore more likely to be resilient
to environmental changes or to exceeding crit-
ical thresholds (Loreau et al. 2001). These gen-
eralized ecological relationships between bio-
diversity and stability have been demonstrat-
ed for both short-term (Loreau et al. 2003) and
geological time scales (Kiessling 2005). For the
current study we use generic richness, even-
ness, and composition of plant communities
as proxies for overall ecosystem persistence
stability, which we dene as the ability to
withstand abiotic perturbations as indicated
by changing isotopic composition.
Richness. Trends in generic richness across
the Tr/J boundary were investigated by stan-
557 TRIASSIC/JURASSIC VEGETATION DYNAMICS
TABLE 3. Precentage representation of leaf genera at Astarteklft from generic abundance matrix (Table 2).
Taxon Bed 1 Bed 1.5 Bed 2 Bed 3 Bed 4 Bed 5 Bed 6 Bed 7 Bed 8
Anomozamites 16.07 32.26 8.14 0.19 27.05 0.10 0.00 0.46 0.00
Baiera 6.25 0.00 0.00 1.33 0.00 0.00 0.00 0.00 0.00
Cladophlebis 0.00 0.00 3.49 0.00 0.00 0.29 39.06 2.55 5.43
Clathropteris 0.00 0.00 1.55 0.00 0.00 0.00 0.00 0.00 0.00
Ctenis 0.00 0.00 0.00 0.00 0.00 0.39 0.00 0.00 0.00
Ctenozamites 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Czekanowskia 2.23 0.00 0.00 0.00 0.00 0.00 0.00 51.16 12.22
Dictyophyllum 0.89 30.65 0.39 0.38 16.44 0.00 8.59 0.00 7.69
Doratophyllum 0.00 0.00 0.00 7.81 0.00 0.00 0.00 0.00 0.00
Elatocladus 0.45 6.45 6.20 0.00 0.11 0.29 14.84 0.00 1.36
Equisetites 10.71 0.00 0.00 0.00 1.14 0.49 1.56 0.00 0.00
Ginkgoites 6.25 0.00 31.01 0.00 0.11 0.00 0.00 24.77 0.00
Hausmannia 0.45 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Lepidopteris 0.45 3.23 4.65 0.00 3.08 0.00 0.00 0.00 0.00
Macrotaeniopteris 0.00 0.00 0.00 0.00 0.00 0.49 0.00 0.00 0.00
Marattia 0.00 0.00 0.00 0.00 0.00 0.00 0.78 0.00 0.00
Matonia 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.71
Neocalamites 0.00 0.00 0.00 0.00 0.00 1.08 0.00 0.00 0.00
Nilssonia 3.13 14.52 0.00 0.57 0.00 0.00 0.00 0.12 0.00
Ontheodendron 0.00 0.00 0.00 0.00 0.00 0.29 0.00 0.00 0.00
Pachypteris 0.00 0.00 0.00 0.00 0.11 0.00 0.00 0.00 0.00
Pagiophyllum 0.00 0.00 0.00 0.00 0.00 0.00 0.78 0.00 0.00
Phlebopteris 0.00 0.00 0.00 0.00 0.00 0.00 0.78 0.00 15.38
Podozamites 27.68 1.61 10.85 86.29 1.48 50.88 0.00 0.00 19.91
Pseudoctenis 0.00 4.84 0.00 0.00 0.11 2.75 0.00 0.00 0.00
Pterophyllum 25.00 4.84 18.99 2.10 50.23 0.00 3.91 0.00 0.00
Ptilozamites 0.00 0.00 0.39 0.00 0.00 0.88 0.78 0.00 0.00
Rhinipteris 0.00 0.00 13.57 0.00 0.00 0.00 0.00 0.00 0.90
Sagenopteris 0.00 0.00 0.00 0.00 0.00 0.00 1.56 0.00 0.00
Schizoneura 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Sphenobaiera 0.00 0.00 0.00 1.14 0.00 0.00 0.00 20.95 34.39
Stachyotaxus 0.45 1.61 0.78 0.00 0.00 40.78 27.34 0.00 0.00
Taeniopteris 0.00 0.00 0.00 0.19 0.11 0.88 0.00 0.00 0.00
Thaumatopteris 0.00 0.00 0.00 0.00 0.00 0.39 0.00 0.00 0.00
dardizing absolute generic richness for all
nine fossil plant beds using rarefaction anal-
ysis. Analytical Rarefaction 1.3 by S. M. Hol-
land was used to estimate rareed leaf generic
richness and 95% condence intervals for sev-
en of the nine fossil plant beds, all of which
had more than 220 specimens (Table 1). In the
case of Beds 1.5 and 6, which had less than 220
specimens (62 and 128, respectively), their re-
spective richness values were forward-pro-
jected to 220 specimens by tting the most sta-
tistically appropriate curve to their respective
rarefaction curves: logarithmic in the case of
Bed 1.5 (R
2
0.99) and power in the case of
Bed 6 (R
2
0.99). Generic richness reported
for these two plant beds should therefore be
considered preliminary. Further collecting
will be undertaken to test these preliminary
richness estimates.
Taphonomic studies on modern leaf litter
demonstrate that multiple samples (four) of
temperate leaf litter capture on average 85%of
the standing species richness of the source
vegetation, whereas subtropical and tropical
leaf litter typically underestimates standing
richness by four and eight times, respectively,
owing to high spatial heterogeneity (Burnham
1993). We have addressed this important po-
tential bias by estimating within-bed hetero-
geneity following the methods of Burnham
(1993), which are interpreted to reect the de-
gree of ecological heterogeneity in the source
vegetation, using Sorensens Coefcient of
Similarity (Sorensens Index, SI; 2C/[A B]),
where C is the number of species in common
between two samples and A and B are the to-
tal number of species in each of the two sam-
ples (Burnham 1993). A mean Sorensens In-
dex was calculated for Beds 1, 2, 3, 4, 5, 7, and
8 from six pairwise comparisons between dif-
558 JENNIFER C. MCELWAIN ET AL.
FIGURE 4. Standing species richness (A), number of species extinctions expressed as a proportion of standing spe-
cies richness (B), species originations (C), range-through species (D), and number of singletons (species occurring
in one fossil plant bed only) expressed as a proportion of A (E). AE calculated from presence absence data of Harris
(1937) for Astarteklft (Appendix 2). See text for more details on methods. Phases 1 to 4 represent distinct evolu-
tionary/ecological phases in the vegetation of East Greenland which are described in detail in the text.
ferent collectors individual quarries, and for
Bed 1.5 from two pairwise comparisons. Rich-
ness estimates derived from forest leaf litter
with Sorensens Indices of less than 50% can
underestimate that of source vegetation by as
much as three times, whereas leaf litter sam-
ples with mean SIs greater than 50% under-
estimate source vegetation richness by only
1.4 times (Burnham 1993).
For comparison we have also estimated
standing species richness (Fig. 4A), species
origination/immigration (Fig. 4C), species ex-
tinction/emigration (Fig. 4B), and range-
through species (Fig. 4D) from Harriss (1937)
original presence/absence data set for Astar-
teklft following the methods of Foote (2000)
and elaborated on for application to paleobo-
tanical specimens by Wilf and Johnson (2004).
This analysis excludes singletons (species oc-
curring in one fossil plant bed only) and as-
sumes the presence of taxa in a plant bed if it
occurs in the beds both immediately above
and below, i.e., with range-through occurrenc-
es (Appendix 2).
Evenness. Changes in the patterns of dom-
inance versus evenness of the vegetation span-
ning the Tr/J boundary were assessed from
analyses of the relative abundance matrix ob-
tained from the census-collected fossil plant
beds of Astarteklft using Shannons Diver-
sity (H) divided by the logarithm of number
of taxa, using PAST Version 1.33. Specical-
ly, we were interested in addressing whether
there was any evidence for ecosystem insta-
bility, as indicated by a decrease in evenness,
prior to the Tr/J boundary. The advantage of
tracking changes in evenness in addition to
richness is that evenness can be accurately es-
timated from the fossil record irrespective of
sample size (Peters 2004).
Results
Biodiversity Richness. There is an 85% de-
cline in standing species richness of plant
559 TRIASSIC/JURASSIC VEGETATION DYNAMICS
FIGURE 5. Rarefaction curves of summed Triassic (Beds
1, 1.5, 2, 3, 4, 5) and Jurassic (Beds 6, 7, 8) leaf generic
richness versus number of macrofossil genera. Phases 1
to 4 represent distinct evolutionary/ecological phases in
the vegetation of East Greenland, which are described
in detail in the text.
FIGURE 6. Rarefaction curves of leaf generic richness
versus number of macrofossil specimens for each of the
nine fossil plant beds at Astarteklft normalized for 220
specimens per bed with the exception of Beds 1.5 and 6
(see Fig. 4).
communities in the Astarteklft region
throughout the Rhaetian with minimumlevels
coinciding with the Tr/J boundary and the
most negative carbon isotope values (Figs. 1,
4A). This marked decline in standing species
richness is attributable to a combination of el-
evated levels of species extinction at and im-
mediately prior to the Tr/J boundary (70% in
Bed 4 and 80% in Bed 5) (Fig. 4B), depressed
levels of species originations (Fig. 4C), and a
sharp decline in the number of range-through
taxa (Fig. 4D), all initiated prior to the depo-
sition of Bed 3 and continuing through the
Rhaetian. At the species level, therefore, spe-
cies turnover at this locality appears to have
increased sharply at the onset of the negative
carbon isotope excursion and remained excep-
tionally high through the latest Rhaetian with
peak proportional extinction coinciding with
the Tr/J boundary.
A comparison of lumped Rhaetian and Het-
tangian rareed leaf generic richness indicates
no signicant differences (Fig. 5), suggesting
that, at the generic level at least, recovery of
Jurassic plant communities following Tr/J
turnover must have occurred relatively rap-
idly (within a few million years). When the
data are broken down into individual beds, a
clearer pattern of changes in generic diversity
emerges (Fig. 6). Beds 1, 1.5, and 2, the oldest
Triassic fossil plant beds, contain the richest
macrofossil assemblages at Astarteklft, with
on average 12 to 13 genera each. A marked
35% loss in generic richness occurs in Beds
3 and 4 (with 7.2 and 8.3 leaf genera, respec-
tively). For Bed 3 in particular, the diversity
loss coincides with the most positive stable
carbon isotope values recorded for the whole
succession. Bed 5, which marks the transition
zone between the Lepidopteris zone of the Tri-
assic and Thaumatopteris zone of the Jurassic,
is richer than Beds 3 and 4, with rarefaction
results indicating 10.9 leaf genera. However,
Bed 5 groups with Beds 3 and 4 in demon-
strating the lowest generic richness among all
nine fossil plant beds analyzed. A conserva-
tive estimate suggests an average generic loss
of 17% between the oldest and youngest (in-
cluding the boundary) Triassic fossil plant
beds. Importantly, the loss occurs in both ge-
nus and species richness, and prior to onset of
the negative carbon isotope excursion but co-
incident with an initial 250 to 500 ppmV rise
in atmospheric CO
2
concentration and pre-
sumed warming of global temperatures (Fig.
1). This suggests that profound changes in
richness and stability of these plant commu-
nities were in place before maximum environ-
mental change occurred at the Tr/J boundary.
The lowest Jurassic assemblage (Bed 6),
which represents the only coal, has an unex-
pectedly high rareed generic richness of 14.5.
560 JENNIFER C. MCELWAIN ET AL.
FIGURE 7. Stable carbon isotopic record from fossil wood (A) compared with changes in rareed leaf generic rich-
ness (B), evenness (which is opposite to ecological dominance) (C), and ecological composition (D), calculated from
relative abundance data derived from a total of 4303 macrofossil leaf specimens from Astarteklft.
It must be noted that this value was extrapo-
lated from 120 to 220 individuals on the basis
of the relationship between rareed generic
richness and 120 individuals. However, even if
further collection of fossil plant specimens
from this bed demonstrates that the true ge-
neric diversity has already been captured
with 120 specimens, a raw diversity measure
of 12 (for 120 individuals) is still higher than
for the underlying three beds. These results
indicate that plant generic richness may have
rebounded rapidly in the Astarteklft region
of Jameson Land (i.e., within a few million
years rather than tens of millions of years) fol-
lowing Tr/J extinctions. As Bed 6 represents
the only coal within the sections at Astar-
teklft, an alternative interpretation of these
results would be that generic richness re-
bounded rapidly among peat-forming plant
communities or that they were unaffected by
events at the Tr/J boundary. We cannot ex-
clude any of these interpretations until a Tri-
assic assemblage with a similar depositional
environment has been identied, for compar-
ison with Bed 6.
Considering richness in isolation from other
measures of biodiversity, these data suggest that
ecosystem instability, as indicated by loss in
plant primary productivity via emigration and
regional extinction, was in place before the Tr/
J boundary. Although generic diversity may
have rebounded rapidly in the Jurassic (Figs. 7,
8B), the depleted levels of standing species di-
versity at Astarteklft (Fig. 4A) in both swamp
and non-waterlogged plant communities sug-
gest that species turnover remained high in the
Jurassic. If the communities rebounded from
Tr/J boundary environmental disturbances,
their composition shows little temporal persis-
tence. This pattern is supported by the obser-
vation that Jurassic plant communities are char-
acterized by an extremely high relative propor-
tion of singletons (on average 40%)an order
of magnitude greater than observed among
those in the Rhaetian (on average 4%)sug-
gesting exceptionally high species ux in the
post-boundary interval.
Biodiversity Evenness. The pattern of
change in generic richness from the Triassic to
Jurassic was in part mirrored by observed
shifts in evenness (Fig. 7C). Plant community
composition was extremely even in the oldest
Triassic plant beds (1, 1.5, and 2) with no sin-
gle taxon dominating. Forest canopy domi-
nants of the levees include Podozamites, Ginkgo,
and Baiera, whereas trees of the oodplain
swamps were represented by Elatocladus,
Stachyotaxus, and to a much lesser extent Po-
dozamites. A rich and even understory of ben-
nettites (Pterophyllum and Anomozamites) oc-
curred in the better-drained areas, whereas
dipteridaceous ferns (Dictyophyllum, Hausman-
nia), cycads (Pseudoctenis and Nilssonia), and
Anomozamites were prevalent in the oodplain
settings. A dramatic decrease in evenness is
evident in Beds 3 and 4. The levee and
561 TRIASSIC/JURASSIC VEGETATION DYNAMICS
FIGURE 8. A, Changes in heterogeneity of the source
vegetation calculated using Sorensens Index (SI) on dif-
ferent quarries from nine fossil plant beds at Astar-
teklft. An SI of 70% indicates that fossil leaf specimens
collected from two different quarries from the same fos-
sil plant bed (and assumed two different samples from
the source vegetation) share 70% of the same taxa. B, Es-
timates of paleolatitude and inferred paleotemperature
based on Reess morphocat system (Rees et al. 2000). A
lower mean DCA score indicates that the composition of
the particular fossil plant bed in East Greenland pro-
vides a signal that was typically associated with lower
paleolatitudes throughout the entire Jurassic and there-
fore from assumed higher paleotemperature (Rees et al.
2000).
swampy oodplain environments became
completely dominated by Podozamites (86% of
relative abundance) and Pterophyllum (50%
relative abundance) respectively. This pattern
of increasing ecological dominance by a single
taxon and decreasing evenness continues to
the Tr/J boundary in Bed 5 where Stachyotaxus
septentrionalis became one of the dominant
taxa. Evenness did not rebound to pre-excur-
sion levels until the youngest Hettangian
plant bed (Bed 8, Fig. 7C).
Heterogeneity. Moving from the oldest to
the youngest fossil plant beds of the Triassic,
a slight but insignicant increase in mean Sor-
ensens Index (SI) is observed, indicating a de-
crease in spatial heterogeneity of the vegeta-
tion through the Triassic (Fig. 8). The most un-
expected shift in ecological heterogeneity oc-
curs in Bed 5 at the Tr/J boundary, which has
a lower SI (47 3%) than all other Triassic and
Jurassic plant beds analyzed. If our paleoeco-
logical data provide an accurate reection of
the ecological makeup of the standing vege-
tation, these results suggest that despite de-
creasing evenness the vegetation was slightly
more patchy at the Tr/J boundary than any
other time before or after the extinction event.
The increased patchiness could reect in-
creased isolation of dry-ground communities
to relatively higher and therefore drier topog-
raphy as wet and swampy habitats favoring
wet-loving plant communities expanded. In-
creased ecological importance of the wet-lov-
ing communities in Bed 5 is indicated by in-
creased relative abundances of Stachyotaxus
(from 2% to 40%) and the presence of Neo-
calamites and Ptilozamites (Appendix 3). An al-
ternative explanation is that patchy extinction
among Late Triassic plant communities and
subsequent invasion by different Jurassic
plant taxa resulted in increased spatial hetero-
geneity in the boundary bed. The shift in het-
erogeneity at the boundary is not, however,
statistically signicant and is unlikely to bias
our estimates of generic richness any more
than the other Triassic and Jurassic plant beds,
as the SI is above 50% if standard errors are
taken into account.
Composition. Marked compositional differ-
ences are evident between Triassic and Juras-
sic plant communities in the region of Astar-
teklft. In the Triassic, four genera, Podozami-
tes, Pterophyllum, Anomozamites, and Dictyo-
phyllum make up on average over 75% of the
relative abundance, but collectively these gen-
era constitute less than 10% of the relative
abundance in Jurassic communities. Similarly,
four taxa (Czekanowskia, Sphenobaiera, Ginkgo
and Cladophlebis) that collectively dominate Ju-
rassic plant communities (89% of relative
abundance) were only very minor compo-
nents of Triassic ecosystems, each contribut-
ing less than 5% of the relative abundance. Re-
sults from detrended correspondence analysis
indicate that the age of the fossil plant assem-
blage exerts a stronger control on composition
than does depositional environment (Fig. 9).
These results suggest large-scale turnover of
the ecological dominants after the Tr/J
boundary, that cannot be explained by taph-
onomic differences between Triassic (all sheet
splay) and Jurassic (swamp and abandoned
channel) fossil plant assemblages. For in-
stance, the oral composition of Bed 1 from
South Tancrediaklft groups more closely
with the same-aged (Triassic) plant beds from
Astarteklft than with assemblages from the
same depositional environment (Fig. 9).
562 JENNIFER C. MCELWAIN ET AL.
FIGURE 9. Scatter plot results of detrended correspon-
dence analysis (DCA) of fossil plant occurrence data
from Astarteklft compared with preliminary occur-
rence data collected from South Tancrediaklft (Tan.).
Triassic fossil plants beds are in italics and Jurassic fos-
sil plant beds are underlined. Note that Bed 1 at Tancre-
diaklft groups more closely with similarly aged fossil
plant beds from Astarteklft (despite differences in
their depositional setting) than it does with Jurassic
aged fossil plant beds at Astarteklft from the same de-
positional setting (abandoned channels). Axis 1, eigen-
value 0.4462; Axis 2, eigenvalue 0.3563.
The rate of compositional change at Astar-
teklft is reected by a stepped change in the
rst eigenvector (axis 1 using DCA) throughout
the Rhaetian and a steady recovery after Tr/J
extinctions (Fig. 7D). Importantly, the exclusion
of genera that occur in only one bed from our
DCA does not change the observed pattern of
compositional change (data not shown). DCA
also reveals that the Tr/J boundary bed (Bed 5)
is the most compositionally distinct of all nine
fossil plant beds analyzed (Fig. 7D). Relative
abundance data from Bed 5 indicate that vege-
tation was co-dominated by Podozamites (50%)
and by Sachyotaxus (40%), a rare component of
the vegetation in the oldest Rhaetian beds, with
an average relative abundance of 2%. The
oodplain ground cover and perhaps subcan-
opy, where present, were dominated almost ex-
clusively by Pseudoctenis with rare occurrences
of Neocalamites, Ptilozamites, and Taeniopteris.
Other paleoecological studies have shown that
such a oristic composition is usually associated
with permanently or frequently ooded envi-
ronments (Archangelsky et al. 1995; Howe and
Cantrill 2001).
These ecological changes parallel those ob-
served for both richness and evenness, indi-
cating that marked changes in plant commu-
nities were initiated before the Tr/J boundary
and
13
C excursion. Together the results more
strongly support a gradual or pulsed trigger-
ing mechanism for Tr/J boundary biodiver-
sity loss rather than a single catastrophic event
coinciding with peak negative isotopic values
used to dene the boundary globally. Long-
term environmental change appears to have
resulted in a major change in the dominance-
diversity structure of plant communities prior
to and coincident with the Tr/J boundary at
Astarteklft.
Ecological Selectivity of Tr/J Extinction
Comparison of the relative abundance distri-
butions of summed Rhaetian and Hettangian
macrofossil plant beds (Fig. 10) indicates that
several key ecological and biological traits are
associated with increased extinction risk at the
Tr/J boundary. These include ecological rarity
and complex reproductive biology, two traits
that are predicted to increase extinction risk
alone or synergistically according to meta-pop-
ulation theory (Gaston 1994; Lawton et al. 1994;
McKinney 1997), and large leaf size, a trait pre-
dicted to be disadvantageous by ecophysiolog-
ical modeling under a Tr/J global warming sce-
nario (McElwain et al. 1999). At the generic
level, extinctions recorded at Astarteklft and in
the Jameson Land region occurred predomi-
nantly among (1) rare genera (dened here as
genera with relative abundances in the last
quartile of the rank abundances, following Gas-
ton, 1994) including dipteridaceous ferns Clath-
ropteris and Hausmannia, the bennettite ower
Weltrichia, and foliage genera Anthrophyopsis (in-
certae sedis, ?Cycadales), Pachypteris (Corystos-
permales), and Macrotaeniopteris (incertae sedis,
?Cycadales); (2) genera with extremely large
leaves (Anthrophyopsis, Clathropteris, Hausmannia,
Macrotaeniopteris); and (3) those genera, judging
from their reproductive anatomy, that likely
would have required specialist insect vectors for
pollination (Wielandiella and Weltrichia).
Physiognomic Selectivity of Tr/J Extinctions.
The selective regional extinction of rare large-
leaved Triassic taxa such as Anthrophyopsis
(20 100 cm), Clathropteris (30 40 cm),
Hausmannia (20 cm in width), and Macro-
taeniopteris (20 50 cm) is consistent with
model expectations based on leaf physiologi-
563 TRIASSIC/JURASSIC VEGETATION DYNAMICS
FIGURE 10. Rank-order abundance plots for summed Triassic versus Jurassic fossil leaf and reproductive genera cal-
culated from 4303 macrofossil specimens collected fromAstarteklft, East Greenland. Dotted line indicates the four quar-
tile (to the right) of the data, which are taxa considered ecologically rare using the denition of Gaston (1994).
cal responses to increased atmospheric CO
2
concentration and mean summer temperature.
McElwain et al. (1999) modeled leaf temper-
ature responses to a fourfold increase in at-
mospheric CO
2
and a 34C global tempera-
ture rise across the Tr/J boundary. They pre-
dicted that large-leaved taxa would become
more dissected and/or reduced in size in or-
der to avoid exceeding lethal leaf temperature
limits under a 5C global warming. Recent pa-
leoclimatic modeling shows that a fourfold in-
crease in atmospheric CO
2
may have raised lo-
cal summer temperatures in the Jameson Land
Basin by 16C, resulting in mean summer tem-
peratures of 36C (Huynh and Poulsen 2005).
Under such high summer temperatures, leaf
temperatures with average undissected leaf
widths of 3 cm would readily exceed 50
55C, a temperature range considered lethal
for tropical/subtropical woody taxa (Gauslaa
1984; Larcher 1994). The demonstration by
McElwain et al. (1999) of the replacement of
large canopy leaves with progressively small-
er and more dissected leaves throughout the
Late Triassic corroborates expectations based
on ecophysiological modeling of leaf temper-
ature. The results presented here suggest that
lethal leaf temperatures may also have ad-
versely affected large-leaved and presumed
understory or ground cover taxa such as Clath-
ropteris, Hausmannia, Anthrophyopsis, and Mac-
rotaeniopteris.
Reproductive Selectivity of Tr/J Extinctions.
Analysis of Harriss (1937) presence/absence
data set for the entire Jameson Land Basin re-
veals that 11 of 13 known bennettite taxa dis-
564 JENNIFER C. MCELWAIN ET AL.
covered in the Kap Stewart Group fossil plant
beds (Wielandiella and Weltrichia) are known
only from the Lepidopteris zone (Triassic). Our
relative abundance data set (Table 3) indicates
that Pterophyllum and Anomozamites, both ben-
nettite foliage taxa, experienced the second
and third most severe average losses in rela-
tive abundances of all taxa recorded from the
Triassic to Jurassic, respectively: Pterophyllum
declines from 20% to 1.3 %, and Anomozamites
from 17% to 0.1% (Table 3). Together these
data suggest that bennettites were severely re-
duced, in terms of both their taxonomic rich-
ness and ecological importance in Jurassic
compared with Triassic plant communities
across the entire Jameson Land Basin. The
protection of ovules among interseminal
scales and within ower-like reproductive
structures (Delevoryas 1963; Harris 1974) and
the presence of nectaries (Crane 1985; Crepet
1974; Delevoryas 1963) are all indicative of a
specialized pollination syndrome, most likely
involving insects, among bennettites.
For cycads, only Doratophyllum experienced
regional extinction, and Ctenis, Pseudoctenis,
and Nilssonia, all of which had low to rare
abundances in the Triassic, are completely ab-
sent from Jurassic plant beds of Astarteklft.
The majority of modern cycad taxa display
mutualistic relationships with highly effective
specialist insect vectors including beetles,
weevils, and thrips (Oberprieler 1995a,b;
Mound and Terry 2001; Terry et al. 2005). Vol-
atiles released from both male and female
cones attract insect vectors to visit, acquire,
and transport pollen (up to 10,000 pollen
grains per individual) between male and fe-
male cones, thus pollinating ovules (Donald-
son 1997). In return, insect vectors receive a
ready food source (pollen from the male and
pollen droplet from the female), protection,
warmth, and oviposition sites (Terry et al.
2005). If these highly mutualistic relationships
between insects and cycads were in place by
the Mesozoic, then extinction of one of the
partners in the relationship would result in
the relatively rapid demise of the other. Recent
observations of Triassic insect coprolites load-
ed with pollen within a permineralized cycad
cone suggest that such mutualistic relation-
ships may have a very long evolutionary his-
tory (Klavins et al. 2003, 2005). We hypothe-
size therefore that the higher levels of extinc-
tion among entomophilous (including bennet-
tites and cycads) than in anemophilous plants
at the Tr/J boundary were caused by a con-
temporaneous extinction of insect taxa. Alter-
natively, extinction and ecological demise of
these plants could have triggered extinction of
their insect vectors. In the absence of insect
body fossils, or a comprehensive record of in-
sect feeding damage from the Jameson Land
Basin, this hypothesis remains to be tested.
The extinction of Lepidopteris at the Tr/J
boundary may also be due to reproductive
specialization; however, this hypothesis is not
conclusive and requires further investigation.
Peltaspermum, the female reproductive struc-
tures of Lepidopteris, consist of relatively com-
plex radially symmetrical peltate ovuliferous
disks from which hang the ovules (Harris
1932a). The micropyle of the ovule in Lepidop-
teris is extremely elongated and angled at
about 30, an anatomical character that may
indicate adaptation for insect rather than wind
pollination. Lepidopteris pollen does not pro-
vide strong support for an entomophilous re-
productive habit, as it is small, 23 to 40 m in
length (Townrow 1960), and unsculptured
with a single sulcus. However, cycads, which
are entomophilous, also have pollen is this
size range. Additional, but admittedly more
indirect, evidence for relatively complex re-
productive biology in Lepidopteris can be
gleaned from its likely habit, which we inter-
pret as vinelike or liana-like (see Appendix 3).
Vegetation Recovery
Twenty-two percent (ve) of fossil plant
genera observed in Jurassic aged plant beds
were not recorded in Triassic assemblages at
Astarteklft, suggesting that they likely im-
migrated from elsewhere. All of these new
Jurassic immigrants can be classied ecologi-
cally as either rare or of low abundance in
terms of their relative abundance distributions
(Fig. 10). Phlebopteris and Matonia (matonia-
ceous fern), Sagenopteris (Caytoniales), Marat-
tia (marattiaceous fern), and Pagiophyllum, all
new genera to East Greenland in the Jurassic,
have relative abundances of less than 2.5%.
Recovery of the plant communities following
the marked biodiversity loss before and coin-
cident with the Tr/J boundary was therefore
565 TRIASSIC/JURASSIC VEGETATION DYNAMICS
achieved by recruitment of new dominants
from within existing plant communities from
the same region, rather than through origi-
nation at the generic level or via signicant
levels of immigration of exotic taxa. For in-
stance, there is evidence for ecological expan-
sion of Cladophlebis,which increased in relative
abundance from 4% in the Triassic to 39%
in the earliest Jurassic plant bed (Bed 6), and
of Stachyotaxus, which increased in abundance
from 2% to 40% at the Tr/J boundary (Bed
5) and to 27% in Bed 6.
Bed 6 at Astarteklft occurs 14 meters above
the Tr/J boundary, but well within the negative
isotopic excursion (Figs. 1, 3) that characterizes
the boundary interval globally. It is noteworthy
in containing the highest proportion of ther-
mophilous elements, which likely represent im-
migrants from much lower paleolatitudes.
These include Pagiophyllum, which is commonly
associated with the thermophilous pollen taxa
Classopollis (Vakhrameev 1991; Axsmith et al.
2004; McElwain et al. 2005) and Sagenopteris. We
estimated an average Jurassic paleolatitude, as a
qualitative proxy for relative global paleotem-
perature, for all nine fossil plant beds following
the methods of Rees et al. (2000). This method
averages the observed paleolatitudes that each
taxon typically occupied in the whole Jurassic
period, using an extensive biogeographical da-
tabase of Mesozoic fossil plants (Rees et al.
2000). The analysis indicates that the taxonomic
composition of Bed 6 is more typical of much
lower paleolatitudes, thus supporting our sug-
gestion that maximum regional temperatures
for the entire RhaetianHettangian interval like-
ly occurred in the earliest Hettangian, coinci-
dent with Bed 6 (Fig. 8B).
Post Tr/J Boundary Fern Spike?
The earliest Jurassic macrofossil plant as-
semblages (represented by Bed 6) are notable
for their high relative proportion of fern taxa.
Bed 6 contains the highest abundance of fern
taxa compared with any of the other Triassic
or Jurassic fossil plant beds: 49% compared
with a maximum of 31% recorded in the Tri-
assic. The high fern sum in Bed 6 is mainly
due to a high relative abundance of Osmun-
daceae (Cladophlebis denticulata); however,
three other fern families are also present (Dip-
teridaceae, Matoniaceae, Marattiaceae). Field-
work in 20022004 shows that this coal, al-
though thin, is laterally extensive across the
whole of the Jameson Land Basin, suggesting
that conditions favoring peat-forming vege-
tation, such as high precipitation and/or re-
duced clastic input, must have prevailed at
this time (Fig. 3). Higher precipitation or an
invigorated hydrological cycle is an expected
consequence of higher global temperatures,
because of increased vapor holding capacity
of the troposphere (Hay and DeConto 1999)
and increased latent heat transfer from low to
high latitudes (Ufnar et al. 2004).
Peat-forming vegetation and a high relative
proportion of fern (trilete) spores (89%) are re-
corded from tropical paleolatitudes in the New-
ark Basin of North America at the Tr/J bound-
ary (Fowell and Olsen 1993; Olsen et al. 2002).
These coaly beds and the contained fernspike
are unique to the Tr/J boundary interval and
have been interpreted as representing an expan-
sion of opportunistic disaster taxa following
the catastrophic environmental effects of a me-
teorite impact (Fowell and Olsen 1993; Olsen et
al. 2002). It is not unusual for peat-forming veg-
etation to contain a high proportion of fern taxa.
It is intriguing, however, that the only coal de-
posits in both basins occur within the negative
isotopic anomaly that characterizes the Tr/J
boundary globally. Further work is now re-
quired to determine whether these high fern
compositions are merely a taphonomic artifact
or whether they signify a truly unique ecologi-
cal and/or climatic event across the Northern
Hemisphere at the Tr/J boundary and in the
earliest Jurassic.
Tr/J Vegetation Dynamics at Astarteklft
Our paleoecological results enable subdi-
vision of the Tr/J vegetation dynamics at As-
tarteklft into four distinct phases. In phase 1,
prior to the negative isotopic excursion and
CO
2
-induced global warming (Fig. 1), the Tri-
assic vegetation of the Astarteklft region was
a rich and heterogeneous broad-leaved, gym-
nosperm-dominated forest. Agathis-dominat-
ed forests of New Zealand and Australia to-
day, with Podocarpus and an understory of the
cycad Lepidozamia hopei, are a good modern
analogue (White 1994). Podozamites, Ginkgo,
Elatocladus, and Baiera most likely co-dominat-
ed the canopy, whereas Pagiophyllum and An-
566 JENNIFER C. MCELWAIN ET AL.
omozamites, and to a lesser extent Nilssonia and
Pseudoctenis, likely formed a sub-canopy. Dip-
teridaceous ferns, Equisetites, and other rare
large-leaved taxa such as Anthrophyopsis and
Macrotaeniopteris made up the ground cover
(Appendix 3). No single taxon dominated the
canopy, sub-canopy, or ground-cover habitats,
and the ora comprised a high percentage of
taxa with complex reproductive cycles requir-
ing animal vectors for fertilization and dis-
persal (cycads and bennettites). If our inter-
pretations are correct, vines were also present
in low to rare abundances (Lepidopteris) (Ap-
pendix 3).
In phase 2, generic richness, evenness, and
standing species richness began to decline
and ecological composition changed marked-
ly. These vegetation changes are indicative of
instability in local plant communities and co-
incide with evidence for deterioration in cli-
matic and atmospheric conditions (McElwain
et al. 1999; Huynh and Poulsen 2005). The can-
opy was rst dominated by Podozamites, and
Stachyotaxus became an additionally impor-
tant element at the boundary. We hypothesize
that a mid-canopy habit in the Triassic was al-
most completely eradicated in the Jameson
Land Basin, owing to high emigration and/or
extinction of erect cycads and bennettites (Ap-
pendix 3). Marked changes in dominance pat-
terns of ground-cover taxa also occurred dur-
ing the Triassic to Jurassic transition as an
abundance of dipteridaceous ferns gave way
to osmundaceous ferns. Phase 3 is character-
ized by low evenness, depressed generic rich-
ness, peak species extinction, and the most
compositionally distinctive vegetation of the
entire RhaetianHettangian interval, as indi-
cated from detrended correspondence analy-
sis. This phase coincides with the rst peak of
the main negative isotopic excursion and max-
imum estimated pCO
2
and global temperature.
The post Tr/J recovery interval, dened
here as phase 4, is characterized by a rebound
of generic richness at Astarteklft; however,
standing species richness for the entire Jame-
son Land Basin remained low throughout this
phase, owing to low species origination and
low numbers of range-through taxa. As Bed 6
is the only coal swamp deposit at Astarteklft,
it is not yet possible to evaluate whether this
apparent rebound in generic richness in Bed 6
is real or a taphonomic artifact. This will be
possible only when a pre-boundary bed from
the same depositional environment has been
identied for direct comparison. Most of the
raw species richness in Hettangian plant beds
at Astarteklft is made up of singletons, sug-
gesting extremely high species turnover dur-
ing the post-boundary interval (Fig. 4E). The
macroecological characteristics of the plant
communities such as evenness and composi-
tion did not fully rebound to pre-excursion
levels until the end of phase 4 as reected in
Beds 7 and 8, which occur 46 m aboveand
likely on the order of 3 to 4 million years af-
terthe Tr/J boundary, indicating a long re-
covery period. It is noteworthy that the nature
and tempo of vegetation dynamics spanning
the Tr/J boundary parallel those observed in
Carboniferous swamp forest spanning the
Westphalian/Stephanian boundary (DiMichele
et al. 1996), hinting that there may be a com-
mon macroecological response of plants to
global climate change.
Signicance of Results in the Broader
Context of Tr/J Boundary Events
Our paleoecological analysis reveals a close
coupling of long-term environmental change
in the latest Rhaetian and marked shifts in the
richness, evenness, and composition of terres-
trial plant communities at Astarteklft. How-
ever, the terrestrial plant communities of the
swamps, oodplains, and dry-ground envi-
ronments show signicant declines in biodi-
versity and shifts in ecological composition
prior to the Tr/J boundary, when maximum
environmental perturbations occurred (Fig.
1). This pattern of more gradual or pulsed bio-
diversity loss and compositional change is in
marked contrast to the tempo of macroecol-
ogical change recorded across the K/T bound-
ary, which was undoubtedly catastrophic
(Wilf and Johnson 2004). A gradual tempo of
extinction and macroecological change prior
to the Tr/J boundary, as recorded here, is not,
however, consistent with a catastrophic causal
mechanism coincident with the boundary,
such as a meteorite impact (Olsen et al. 2002).
The absence of both shocked quartz (Moss-
man et al. 1998) and a suitably aged crater
(Hodych and Dunning 1992), and iridium val-
ues that are more indicative of a mantle than
567 TRIASSIC/JURASSIC VEGETATION DYNAMICS
extraterrestrial origin (Mossman et al. 1998)
also suggest that such a causal mechanism
was rather unlikely.
Global cooling has also been invoked as a
potential extinction mechanism because of an
increased abundance of conifer pollen at the
boundary, which is interpreted as indicative of
climatic cooling (Hubbard and Boulter 2000).
However, the fact that dipteridaceous fern
spores, which today have an exclusively trop-
ical distribution, and Classopollis pollen, a pre-
sumed high-temperature indicator (Vakhra-
meev 1991), are both found in the same pal-
ynological associations as the conifer pollen
strongly argues against a global cooling hy-
pothesis. The role of sea-level changes via
habitat loss in either regional or global Tr/J
marine extinctions has been strongly chal-
lenged (Hesselbo et al. 2004) and lacks a mech-
anistic explanation for gradual extinctions on
land. Our paleoecological analysis most
strongly supports a central role for extinction
mechanisms that operated over hundreds of
thousands to millions of years, initiating en-
vironmental change and instability of plant
communities and perhaps of entire ecosys-
tems. Climatic and environmental change as-
sociated with the emplacement of the Central
Atlantic Magmatic Province (CAMP) is the
most likely candidate. Peak volcanic activity of
CAMP (199.5 0.5 Ma) (Knight et al. 2004;
Marzoli et al. 1999, 2004) is temporally con-
temporaneous with the Tr/J boundary (as de-
ned by the
13
C curve; 199.6 0.3 Ma) and
with minimum negative isotopic values from
organic and inorganic carbon (Palfy 2000).
Ash (1986) suggested that there is little ev-
idence for loss of higher taxonomic ranks of
plants across the Tr/J boundary. Our analysis
in part conrms this, as only one family (Pel-
taspermaceae) that goes extinct in Greenland
is a conrmed global extinction. An absence of
family-level extinction may be an artifact of
taxonomy (Kerp et al. 2006). However, genus-
level oral extinctions at Astarteklft were
also moderate at 17%, despite exceptionally
high species turnover (80%) (Fig. 4B). Using
these data, one could argue that terrestrial
plants do not undergo mass extinction accord-
ing to the strictest denition of the term(in the
sense of Sepkoski, [1981]). In contrast, paleo-
ecological changes at Astarteklft indicate
that terrestrial vegetation in this region likely
underwent major structural and composition-
al change. This pattern of resilience among
higher taxonomic ranks in spite of ecological
upheaval has been observed for marine inver-
tebrates (Droser et al. 1997, 2000). The vege-
tation changes at Astarteklft represent the
second highest level of paleoecological change
of four possible levels developed by Droser et
al. (1997), and far exceed normal background
ecological change. These include likely loss of
the mid-canopy habit, a drastic decline in
abundance of reproductively specialized taxa
(cycads and bennettites) across the entire Ja-
meson Land Basin and a marked change in
plant dominance. Our data suggest that these
changes cannot be explained by taphonomic
differences between Tr/J depositional envi-
ronments (Fig. 9).
We hypothesize, on the basis of evidence for
geographically extensive macrofossil and pal-
ynological turnover (80% of species) across
the entire Jameson Land Basin (Harris 1937;
Pedersen and Lund 1980), that the nature and
magnitude of paleoecological changes record-
ed at Astarteklft reect wider regional veg-
etation change. Changes in the source and/or
structure of plant primary productivity can
have a cascade effect throughout the entire
ecosystem (Vermeij 2004). How exactly these
macroecological changes in the vegetation of
Astarteklft scale up to the whole Jameson
Land region and how they affected the well-
documented faunal extinction at the Tr/J
boundary remain important avenues of future
research.
Conclusions
1. Terrestrial plant biodiversity (richness
and evenness) shows evidence for a gradual
decline at both genus and species levels in the
latest Triassic prior to the Tr/J boundary. This
pattern suggests that terrestrial plant com-
munities at Astarteklft show signs of insta-
bility prior to maximal global climatic and at-
mospheric change. We conclude therefore that
relatively minor changes in CO
2
and regional
temperature had a disproportional negative
effect on the stability of terrestrial plant com-
munities. Alternatively, some as yet undetect-
ed biotic or abiotic factors triggered instability
in advance of the Tr/J boundary.
568 JENNIFER C. MCELWAIN ET AL.
2. Post-Tr/J boundary plant communities are
characterized by an extremely high relative pro-
portion of taxa occurring in one bed only (on
average 40%), compared with 4%in Triassic
plant communities. This indicates exceptionally
high species-level ux in the post-boundary in-
terval. These data, together with a long time de-
lay in the recovery of evenness, suggest a long
post-Tr/J recovery period.
3. Our paleoecological analysis reveals
complete replacement of the dominant Trias-
sic plant taxa with new dominants in Jurassic
plant communities that cannot be explained
simply by taphonomic factors. The Triassic
dominants did not go extinct at the Tr/J
boundary but instead persisted as ecologically
rare components of Jurassic plant communi-
ties. Similarly, the new Jurassic dominants
were recruited from rare occurrences within
Triassic ecosystems.
4. Ecological rarity, large leaf size, and
complex reproductive biology were the pri-
mary ecological/biological traits identied to
increase extinction risk for terrestrial plant
taxa in the Jameson Land Basin at the Tr/J
boundary. All three determinants of extinc-
tion risk match expectations based on modern
meta-population theory and ecophysiological
models.
5. The gradual nature of terrestrial plant
biodiversity decline and compositional
change recorded in East Greenland prior to
the Tr/J boundary does not support a single
catastrophic extinction mechanism coincident
with the boundary as dened by the isotopic
excursion, but rather invokes more gradual
causal agents of plant extinction.
6. Our study of vegetation dynamics span-
ning the Tr/J boundary suggests that the tax-
onomic severity of this extinction was decou-
pled from the ecological severity.
7. If the paleoecological changes at Astar-
teklft were a basin wide phenomenon, as is
highly probable according to preliminary
data, they are likely to have contributed sig-
nicantly to Tr/J vertebrate extinctions by al-
tering the dominant sources of primary pro-
ductivity available to consumers.
Acknowledgments
We thank the National Geographic Society
(7038-01), the Comer Foundation of Science
and Education (#13), the Field Museum, the
Royal Society of London, the United Kingdom
Natural Environment Research Council, and
an EU Marie Curie Excellence Grant (MEXT-
CT-2006-042531) for funding. Special thanks
to E. C. Meeker and to R. H. and P. O. Schnadig
for additional funding support through the
Field Museum Womens Board Field Dream
Program. We thank I. Glasspool and D. Sun-
derlin for eldwork carried out in 2004, which
provided preliminary data on South Tancre-
diaklft. Many thanks to the Danish Polar
Centre, Constable Pynt Airbase staff, and R.
Hines for logistical support with eldwork in
East Greenland. P. Wagner and I. Glasspool
are thanked for comments on earlier versions
of the manuscript. We thank D. Cantrill, W. A.
DiMichele, and one anonymous reviewer for
their extremely thorough and constructive re-
views. This paper is a contribution to Inter-
national Geoscience Programme Project 458.
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Appendix 1
Comparison of different methods of fossil leaf counting on generic relative and rank order abundance in Bed 1,
Astarteklft.
Macrofossil
taxon
Generic leaf/leaf fragment
occurrence
Hybrid method
Quadrat
method
Generic relative abundance (%)
Hybrid method
Quadrat
method
Generic rank order
Hybrid method
Quadrat
method
Pterophyllum 56 36 25.00 27.27 2 1
Podozamites 62 27 27.68 20.45 1 2
Anomozamites 36 26 16.07 19.70 3 3
Ginkgoites 14 11 6.25 8.33 5 4
Baiera 14 10 6.25 7.58 5 5
Equisetites 24 8 10.71 6.06 4 6
Nilssonia 7 7 3.13 5.30 6 7
Dictyophyllum 2 2 0.89 1.52 7 8
Czekanowskia 5 1 2.23 0.76 8 9
Elatocladus 1 1 0.45 0.76 9 9
Hausmannia 1 1 0.45 0.76 9 9
Lepidopteris 1 1 0.45 0.76 9 9
Stachyotaxus 1 1 0.45 0.76 9 9
Appendix 1A
Comparison of different counting methods on standard measures of diversity of fossil leaf taxa in Bed 1, Astar-
teklft.
Hybrid method Quadrat method Hybrid method Quadrat method
Taxa S 13 13 Evenness e
H/S
0.5214 0.5547
Individuals 224 132 Menhinick 0.8686 1.132
Dominance D 0.1859 0.1747 Margalef 2.217 2.458
Shannon H 1.914 1.976 Equitability J 0.7461 0.7702
Simpson 1-D 0.8141 0.8253 Fisher alpha 3.006 3.576
Berger-Parker 0.2768 0.2727
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Appendix 2
Range through species data for Astarteklft from Harris (1937: Table 1).
Note: 1 denotes presence and 0 denotes absence. Shaded gray boxes indicate where fossil plant taxa are assumed to be present given their presence
in bed both above and below. 1? denotes questionable occurences.
Appendix 3
Autecological interpretation of Tr/J genera discussed in the text.
Stachyotaxus Tree habit, wet-loving,
swamp conifer with simi-
lar habit to modern Taxo-
dium distichum.
Tree habit is inferred from taphonomy studies that show that
canopy trees are best represented in the fossil record, and
that a herbaceous or shrubby habit is unknown and rare, re-
spectively, for conifers.
Similarities in overall leaf habit to Taxodium distichum (leaves
shed in long shoots, thin cuticle, leaf mats of Stachyotaxus
common suggestive of a deciduous habit).
Ecological association with common oodplain taxa such as Ano-
mozamites, Dictyophyllum, and Pseudoctenis (Archangelsky et al.
1995) indicated using detrended correspondence analysis (DCA).
Fossil evidence suggesting extremely high seed output by Stach-
yotaxus also supports this ecological interpretation, as this is a
common strategy among modern taxa of frequently ooded
and waterlogged habitats that increases the chances for seed-
ling establishment (Parolin 2001).
Sedimentological evidence that suggests Stachyotaxus occurred
in semi- or permanently ooded oodplains and more abun-
dantly in coal swamps.
Podozamites Tree habit; broad-leaved, like-
ly deciduous; conifer; domi-
nant; occurs in strongest as-
sociation with Pterophyllum
and rarely with wet-loving
taxa (Equisetites) or ood-
plain taxa (such as Pseudoc-
tenis).
A tree habit is likely because leaf mats of Podozamites are com-
mon and parautochthonous depositional environments tend
to favor representation by canopy trees.
Large trunks (2 m in length) are commonly found with Podoz-
amites leaf remains in Hettangian deposits from Romania
(Popa 2000).
Gross leaf morphology similar to some species of modern
broad-leaved conifers such as Agathis (White 1994).
Cuticles thin and leaf mats abundant, suggestive of a deciduous
habit.
Small fragments of coalied and charcoalied wood are com-
mon in beds dominated by Podozamites.
Dictyophyllum Dipteridaceous fern; can with-
stand temporary ooding;
closely associated with Nils-
sonia and Anomozamites
Detrended correspondence analysis of occurrence data for Beds
1 to 5 shows Anomozamites closely associated with Nilssonia,
Stachyotaxus, and Dictyophyllum.
Best represented in Bed 1.5.*
Modern Dipteris ferns are exclusively tropical and are often col-
onizers of open disturbed habitats.
573 TRIASSIC/JURASSIC VEGETATION DYNAMICS
Appendix 2. Extended.
Appendix 3. Continued.
Lepidopteris Vinelike or liana-like habit;
not common; most closely
associated with Elatocladus
A main stem to pinna rachis ratio of 5:1, leaf heterophylly and
thick cuticle, deeply sunken stomata and possible hyda-
thodes, referred to by Harris (1932a) as abnormal stomata;
vein endings that widen toward the leaf margin; and leaf and
stem surfaces covered in blisterlike structures that proba-
bly contained anti-herbivore defense compounds. All of these
anatomical and morphological characteristics are common
among modern angiosperm vines and lianas, which common-
ly display xeromorphic adaptations, as the vining habit
makes it difcult to maintain adequate water supply through-
out the plant body, despite their occurrence in moist/humid
environments (Krings et al. 2003).
Ginkgo Tree habit; adapted to dry
ground rather than tempo-
rary or permanently water-
logged environments; decid-
uous
A tree habit is likely because leaf mats of Ginkgo are common
and parautochthonous depositional environments tend to fa-
vor representation by canopy trees (Gastaldo 2001).
Ginkgo does not occur in the plant beds that have the best rep-
resentation from in situ oodplain communities (i.e., Bed 1.5).
It likely occupied dry ground on the levees, rather than wa-
terlogged environments of the oodplain or swamps.
Pagiophyllum Form genus of scale-leaved co-
nifers
This form genus typically occurs at lower latitudes than the Ja-
meson Land Basin in the Jurassic suggesting that plants
which produced these leaves were warm adapted (Vakhra-
meev 1991; Rees et al. 2000; McElwain et al. 2005).
High-resolution mesofossil studies of the Toarcian have shown
peaks in Pagiophyllum coinciding with peak inferred global
temperatures (McElwain et al. 2005).
Anomozamites Can withstand temporary
ooding, common on ood-
plain in association with
Dictyophyllum and Nilssonia
Detrended correspondence analysis of occurrence data for Beds
1 to 5 show Anomozamites closely associated with Nilssonia,
Stachyotaxus, and Dictyophyllum.
Best represented in Bed 1.5.*
Nilssonia Likely deciduous cycad adapt-
ed to temporary ooded en-
vironments
Best represented in Bed 1.5.*
The plant to which Nilssonia foliage belongs has been interpreted
as a deciduous, frost-resistant, woody cycad that can rapidly shed
leaves in clusters attached to short shoots (similar to that of Gink-
go) (Spicer and Herman 1996) or as individual leaves (Kimura and
Sekido 1975).
Pseudoctenis Cycad adapted to temporary
ooded environments
In situ whole plants have been recorded in oodplain environ-
ments (Archangelsky et al. 1995).
Best represented in Bed 1.5.*
In Bed 5 it has a highly heterogeneous lateral distribution com-
pared with other taxa.
* Judging from very high preservation quality of some of the largest leaves/fronds recovered from the entire succession, Bed 1.5 most likely indicates
good representation from in situ oodplain communities.